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THE    ANIMAL 
PARASITES   OF    MAN 


BY 

H.    B.    FANTHAM,    M.A.Cantab,    D.Sc.Lond. 

Lecturer  on   Parasitology,   Liverpool   School  of  Tropical  Medicine ;    iSectionctl  Editor   in 
Protozoology^  "  Tropical  Diseases  Bulletin,^'  Londofi,  atc.^         •"''', 

J.    W.    W.    STEPHENS,    M.D.Cantab.,    D.P.H. 

Sir  Alfred  [o7ies  Professor  of  Tropical  Medicine^   Liverpool  University^  etc. 

AND 

F.  V.  THEOBALD,  M.A.Cantab.,  F.E.S.,  Hon.  F.R.H.S. 

Professor  of  Agricultural  Zoology,  London  University  ;     Vice- Principal  and  Zoologist  of  the 

South-eastern  Agricultural  College  ;    Mary  Kingsley  Medallist  ;     Grande  Medaille  Geoffroy 

St.  Hilaire,  Soc.  Nat.  d'' Accliui.  de  France^  etc. 


PARTLY    ADAPTED    FROM 

Di.  Max  Braun's  *'  Die  Tierischen  Parasiten   des  Menschen  "  (4th  Edition,  1908)  and  an 
Appendix  by  Dr.  Otto  Seifert. 


NEW     YORK 

WILLIAM      WOOD      AND      COMPANY 

MCMXX. 


LIBRARY 

.UNIVERSITY  OF  CALIFORNL\ 
DAVT.S 


N 


PREFACE  111 


PREFACE. 

The  English  edition  of  Braun's  ''  Die  Tierischen  Parasiten  des 
Menschen,"  produced  in  1906,  being  out  of  print,  the  pubhshers 
decided  to  issue  another  edition  based  on  the  translation  of  Brain's 
fourth  German  edition,  which  appeared  in  1908,  to  which  had  been 
added  an  appendix,  by  Dr.  Otto  Seifert  on  Treatment,  etc/    •       ^ .'. 

When  the  work  was  considered  with  a  view  to  A  hevv  edition, 
it  was  found  that  a  vast  amount  of  new  matter  had  to  be  incorporated, 
numerous  alterations  essential  for  bringing  it  up  to  date  were  necessi- 
tated, and  many  omissions  were  inevitable.  The  result  is  that  parts  ol 
the  book  have  been  rewritten,  and,  apart  from  early  historical  references, 
the  work  of  Braun  has  disappeared.  This  is  more  particularly  the  case 
with  the  Protozoa  section  of  the  present  work.  The  numerous  addi- 
tions, due  to  the  great  output  of  scientific  literature  and  other  delays  in 
publication,  have  led  to  the  book  being  somewhat  less  homogeneous 
than  we  desired,  and  have  necessitated  the  use  of  appendices  to 
allow  of  the  presentation  of  new  facts  only  recently  ascertained. 
Many  new  illustrations  have  been  added  or  substituted  for  older,  less 
detailed  ones.  Some  of  these  new  figures  w^ere  drawn  specially  for 
this  book* 

The  first  section,  on  the  Protozoa,  has  been  wTitten  by  Dr.  Fantham, 
there  being  little  of  the  original  text  left  except  parts  of  the  historical 
portions,  and  thus  the  section  on  Protozoa  must  be  considered  as 
new.  The  second  section,  on  Worms  (except  the  Acanthocephala, 
Gordiidae  and  Hirudinea),  has  been  remodelled  by  Professor  Stephens 
to  such  an  extent  that  this,  too,  must  not  be  looked  upon  as  a 
translation  of  Braun's  book.  With  regard  to  the  Arthropoda, 
much  remains  as  in  the  last  English  edition,  but  some  new  matter 
added  by  Braun  in  his  fourth  German  edition  is  included,  and  much 
new  matter  by  Mr.  Theobald  has  been  incorporated.  As  regards 
the  Appendix  by  Dr.  Seifert,  the  first  section  has  been  remodelled, 
but  the  sections  on  the  Helminthes  and  the  Arthropoda  are  practically 
translations  of  the  original. 

The  authors  desire  to  express  their  thanks  to  Miss  A.  Porter,  D.Sc, 
J.  P.  Sharpies,  Esq.,  B.A.,  M.R.C.S.,  and  H.  F.  Carter,  Esq.,  F.E.S., 
for  valuable  help.     They  also  wish  to  thank  the  authors,  editors,  and 


IV  THE   ANIMAL  PARASITES   OF   MAN 

publishers  of  several  manuals  and  journals  for  their  courtesy  in 
allowing  the  reproduction  of  certain  of  their  illustrations.  In  this 
connection  mention  must  be  made  more  particularly  of  Professor 
Castellani,  Dr.  Chalmers,  Professor  Doflein,  Dr.  Leiper,  the  late 
Professor  M  inch  in.  Professor  Nuttall,  Dr.  Wenyon,  Mr.  Edw.  Arnold, 
Messrs.  Bailliere,  Tindall  and  Cox,  Messrs.  Black,  Messrs.  Cassell, 
Dr.  Gustav  Fischer,  Messrs.  Heinemann,  the  Cambridge  University 
Press,  the  Editors  of  the  Annals  of  Tropical  Medicine  and  Parasitology, 
the  Editors  of  the  Journal  of  Experimental  Medicine,  and  the  Editor 
of  the  Tropical  Diseases  Bnlletin. 

H.  B.  F. 
J.  W.  W.  S. 
December,  J915.  F.  V.  T. 


CONTENTS 


CONTENTS, 


PAGE 

PREFACE   ...            ...            ...             ...            ...             ...             ...  ...            ...  Hi 

ERRATA ^ xxxii 

ON   PARASITES    IN    GENERAL i 

Occasional  and  Permanfent  Parasitism               ...             ...             ...  ...              ...  i 

Entozoa,  Endoparasites,  Helminthes,  Turbellaria          ...             ...  ...              ...  2 

Hermaphroditism                  ...              ...              ...              ...              ...  ...              ..  4 

Fertility  ot  Parasites            ...              ...             ...             ...              ...  ...             ...  5 

Transmigrations    ...             ...              ...             ...             ...              ...  ...             ...  5 

Commensals,  Mutualists      ...             ...              ...             ...             ...  ...             ...  6 

Incidental  and  Pseudo-parasites         ...             ...             ...               ..  ...             ...  6 

The  Influence  of  Parasites  on  the  Host            ...             ...             ...  ...             ...  8 

Origin  of  Parasites                ...              ...              ...              ...              ...  ...              ...  10 

Derivation  of  Parasites         ...             ...             ...             ...             ...  ...             ...  19 

Change  of  Host    ...             ...              ...             ...             ...              ...  ...             ...  20 

Literature              ...             ...             ...             ...             ...             ...  ...             ...  22 


THE  ANIMAL   PARASITES   OF   MAN         25 

A.  Protozoa       ...  ...  ...  ...  ...  ...  ...      25 


Classification  of  the  Protozoa. 


Class  I. 

Sarcodina  ... 

Order. 

Amcebina 

Foraminifera 

Heliozoa     . . , 

Radiolaria 

Class    II. 

Mastigophora 

III. 

Sporozoa 

Sub-class 

I.     Telosporidia 

Order. 

Gregarinida 

Coccidiidea 

Hamosporidia 

Sub-class 

2.     Neosporidia 

Order. 

Myxosporidia 

Microsporidia 

Sarcosporidia 

Haplosporidia 

Class  IV. 

Infusoria 

V. 

SUCTORIA 

27 

27 
27 
27 
28 
28 
28 
28 
28 
28 
28 
28 
28 
28 
28 
29 
29 
29 


VI  THE  ANIMAL   PARASITES   OF   MAN 

The  Animal  Parasites  of  Man — contd, 

ProtOZOSi— con  ^d.  PAGE 

Class  I.     Sarcodina,  Biltschli,  1882  ...  ...  ...  ...  ...  29 

Order.     Amcebina,  Ehrenberg   ...  ...  ...  ...  ...  ...  29 

A.  Human  Intestinal  Amoeb£e  ...  ...  ...  ...  ...  29 

Entamceha  coli,  Losch,  1875,  emend.  Schaudinn,  1903  ..  32 

Entamoeba  histolytica,  Schaudinn,  1 903    ...  ...  ...  34 

Enta/nceba  te(rage7ta,  WiQxecV,  1907...  ...  ...  38 

Noc's  Entamoeba,  1909  ...  ...  ...  .  .  41 

Entamceba  buccalis,  Prowazek,  1904  ...  ...  ...  43 

Entatnceba  undulans,  QsL^itWaxviy  i()0^       ...  ...  ...  43 

Entamoeba  kartulisi,  Doflein,  1901  ...  ...  ...  44 

A7no;ba  gingivalis,  A.  buccalis,  A.  dentalis  ...  ...  44 

Genus.     Paramceba,  Schaudinn,  1896  ...  ...  ...  44 

Paramceba  (Craigia)  hominis,  Craig,  1906  ...  ...  45 

B.  Amoebae  from  other  Organs         ...  ...  ...  ...  ..  45 

Entamoeba pult?ionalis,  Artault,  1898        ...  ...  ...  45 

Amceba  urogeititalis ,  Baelz,  1883  ...  ...  ...  45 

Aviceba  miurai,  Ijima,  1898        ...  ...  ...  ...  46 

Appendix:  "  Rbizopods  in  Poliomyelitis  Acuta"  ...  ...  ...  46 

Order.     Foraminifera,  d'Orbigny  ...  ...  ...  ...  ...  47 

Sub-order.     Monothalamia  (Tc-taceous  Amoebse)  ...  ...  ...  ...  47 

Genus.     Chlamydophrys,  Cienkowski,  1876    ...  ...  ...  47 

Chlamydophrys  enchelys,  Ehrenberg         ...  ...  ...  47 

Leydenia geinmipara,  Schaudinn,  1896     ...  ...  ..  49 

Class  II.     Mastigophora,  Diesing  ...  ...  ...  ...  ...  50 

Sub-class.     Flagellata,  Cohn  emend.  Biitschli  ...  ...  ...  ...  50 

Order.     Polymastigina,  Blochmann  ...  ...  ...  ...  ...  52 

Genus.     Trichomonas,  Donne,  1837  ...  ...  ...  52 

Trichomonas  vaginalis,  Donne  ...  ...  ...  ...  52 

Trichomonas  intestinalis,  R.  Leuckart,  1 879  =   Trichomonas 

hominis,  Davaine,  1854   ...  ...  ...  ...  54 

Genus.      Tetramitus,  Perty,  1852      ...  ...  ...  ...  57 

letratnitus  mesnili,^Qnyox\y  1910  ...  ...  ...  57 

Genus.     Lamblia,  R.  Blanchard,  1888  ...  ...  ...  57 

Lamblia  intestinalis,  Lambl,  1 8 59  ...  ...  ...  57 

Order.     Protomonadina,  Blochmann        ...  ...  ...  ...  ...  60 

Family.      Cercomonadida,  Yieni  evaerxA. 'RviischW    ...  ...  ...  61 

Genus.      Cercomonas,  Dujardin  emend.  Biitschli  ...  ...  61 

Cercomonas  hominiSf'Da.vdXnG,  1 854  ...  ...  ...  61 

Monas pyophila,  R.  Blanchard,  1895         •■•  •••  •••  ^2 

Family.     Bodonida,  Biitschli      ...  ...  ...  ...  ...  63 

Genus.     Prowazekia,  Hartmann  and  Chagas,  19 10       ...  ...  63 

Prowazekia  urinaria,  Hassall,  1859  ...  ...  ...  63 

Prowazekia  asiatica,  C2i5\.Q\\2iX\i  Siwd  OmXmex?,,  1910  ...  65 

Prowazekia  favajiensis,Y\n,  igi2  ...  ...  ...  66 

Prowazekia  crttzi,  Hartmann  and  Chagas,  igio      ...  ...  66 

Prowazekia  weinbergi,  MsLthis  and  heger,  igio      ...  ...  66 

Prowazekia  parva,  Nagler,  1910  ...  ...  ...  66 

Family,      Trypanoso?nidcB,  Doflein  ...  ...  ...  ...  66 

Genus.      Trypanosoma,  Gruby,  1843  •••  •••  •••  ^7 

Historical  ...  ...  ..  ...  ...  67 

General  ...  ...  ...  ...  ...  ...  69 

Morphology  ...  ...  ...  ...  ...  70 

Trypanosoma  gainbiense,  Dutton,  1902     ...  ...  ...  72 

Trypanoso?na  nigeriense,  Macfie,  1913      ...  ...  ...  76 


CONTENTS  Vii 

The  Animal  Parasites  of  Man — contd. 

Protozoa.— con^d.  page 

Trypanosoma  rhodesiense,  Stephens  and  P'antham,  19 lo       ...  76 

General  Note   on  Trypanosomes  with  Posterior  Nuclei  83 

Trypanosoma  crt4zi,  C\\2ig2i%,  1 909              ...  ...  ...  83 

Trypanosotna  lewisi,  Kent,  1 88 1                ...  ...  ...  88 

Trypanosoma  brucei,  Plimmer  and  Bradford,  1899...  ...  93 

Trypanosoma  evansi^  Steel,  1885                ...  ...  ...  95 

Trypanosoma  equinum^V ogQs,  igoi         ...  ...  ...  96 

Trypanosofna  equiperdtwi^  Doflein,  1901  ...  ...  ...  97 

Trypanosoma  theileri^  Bruce,  1902            ...  ...  ...  98 

Trypanosoma  hippicum^  Darling,  1910      ...  ...  ...  98 

Endotrypamim  schaudinni,  Mesnil  and  Brimont,  1908  ...  99 

Trypanosoma  boy  let,  Lafont,  1912              ...  ...  ...  99 

Monomorphic  Trypanosomes              ...             ...  ...  ...  99 

Trypanosoma  vivax,  7.\Qvi\2iX\v\,  1905          ...  ...  ...  99 

Trypanosoma  caprce,  Kleine,  1910              ...  ...  ...  lOO 

Trypanosoma  con^olenscy  Broden,  1904     ...  ...  ...  100 

Trypanosoma  simice,'Qx\xcCi  1912               ...  ...  ...  lOO 

Trypanosoma  uniforme.,  Bruce,  1910         ...  ...  ...  loi 

General  Note  on  Development  of  Trypanosomes  in  Glossina  10 1 

Adaptation  of  Trypanosomes                ...  ...  ...  lOl 

Genus.     Herpetomonas,  Saville  Kent,  1881      ...  ...  ...  102 

Genus.     Crithidia,  Leger,  1902,  emend.  Patton,  1908...  ...  104 

Genus.     Leishmania,  Ross,  1903       ...             ...  ...  ...  104 

Zm/^wa«2'a  «?<?«cz/a««,  La veran  and  Mesnil,  1903  ...  ...  105 

Leiskma7tia  tropica,  Wng\it^  1^01^              ...  ...  ..  107 

Leishmania  infantum,  Nicolle,  1908        ...  ...  ...  109 

Genus.     Histoplasma,  Darling,  igo6                ...  ...  ...  112 

Genus.      T^^xo/Zaj-wat,  Nicolle  and  Manceaux,  1908  ...  ...  112 

The  SpirocH/^tes     ...            ...            ...            ...            ...  ...  ...  114 

The  Spirochaetes  of  the  Blood             ...             ...  ...  ...  116 

6'/zV^c/z^/a  ^w//^«?,  Novy  and  Knapp,  1906  ...  ...  116 

*  Spirochata   gallinaj-um,    Stephens    and   Christophers,    1905 

(=  Spirochata  marchouxi,  Nuttall,  1905)  ...  ...  119 

Spirochcsta  recm-ren/is,  l^ehtxt,  i?)"]/^          ...  ...  ...  120 

SpirochcBta  rossii,  Nuttall,  1908                  ...  ...  ...  122 

Spiroc hteta  novyi,  ScheWdick,  igoj             ...  ...  ...  122 

SpirochcBta  carter i,  Mackie  and  Manson,  1907  ...  ...  122 

SpirochcBta  berbera,  Sergent  and  Foley,  1910  ...  ...  122 

Other  Human  Spirochaetes                   ...              ...  ...  ...  122 

Some  Animal  Spirochoetes  ...             ...             ...  ...  ...  122 

Treponemata             ...            ...            ...            ...            ...  ...  ...  124 

Treponema  pallidum,  Schaudinn,  1905     ...  ...  ...  124 

Treponema pertenue,  Castellani,  1905        ...  ...  ...  127 

Class  III.     Sporozoa,  Leuckart,  1879            ...             ...             ...  ...  ...  128 

Sub-class.     Telosporidia,  Schaudinn            ...             ...             ...  ...  ...  129 

Order.     Crif^a^-zV^/^a;,  Aime  Schneider  emend.  Doflein         ...  ...  ...  129 

Order.     Coccidiidea      ...             ...             ...             ...             ...  ...  ...  135 

Genus.     ^?///^rm,  Aime  Schneider,  1875         •••  •••  •••  '4^ 

.£"/»?^r2a  az/mw,  Silvestrini  and  Rivolta   ...  ...  ...  142 

Eimeria  stied(B,\Jvci'\^xtvzxiX\,  \%(i^             ...  ...  ...  145 

(a)  Human  Hepatic  Coccidiosis         ...  ...  ...  148 

(b)  Human  Intestinal  Coccidiosis      ...  ...  ...  148 

{c)  Doubtful  Cases                ...             ...  ...  ...  149 


viii  THE  ANIMAL   PARASITES   OF   MAN 

The  Animal  Parasites  of  Man — contd. 

Protozoa.— confd.  pa(;e 

Genus.     Isospora^  Aime  Schneider,  i88i         ...  ...  ..  I49 

Isospora  bigemina^  Stiles,  1891  ...             ...  ...  •••  149 

Doubtful  Species  ...             ...             ...             ...  •••  •••  150 

Order,     ^^woi/^jr/i/za,  Danilewsky  emend.  Schaudinn        ...  ...  ...  151 

The  Malarial  Parasites  of  Man        ...  ...  ...  155 

Development  of  the  Malarial  Parasites  of  Man  ...  159 

The  Species  of  the  Malarial  Parasites  of  Man...  ...  ...  164 

Plasmodium  vivax,  Grassi  and  Feletti,  1890  ...  ...  164 

PlasfTiodium  malaria: ,  Laveran  ...              ...  ...  ...  166 

Laverania  malarice,  Grassi  and  Feletti,  1890  {=  Plasmodium 

falciparum,  ^€\c\\,i?>9'j)                ...  ...  •••  167 

/•/ai^w^^/Mw  7^/zV/Mw,  Sergent,  1907  (in  birds)  ...  >..  170 

Cultivation  of  Malarial  Parasites        ..  ...  ...  170 

Differential  Characters  of  the  Human  Malarial  Parasites  171 

P'amily.     Piroplasmidce,  Franca,  1909      ...             ...  .••  ...  172 

Genus.     Babesia,  Starcovici,  1893    ...             ...  ...  ...  174 

Genus.      Ty^^zV^rza,  Bettencourt,  Fran9a  and  Borges,  1907  ...  178 

Theileria parva,  Theiler,  1903  ...              ...  ...  ...  178 

Iheileria  mutans,  Theiler,  1907            \     ...  ...  ...  1 80 

Genus.     Anaplasj/ia,  Theiler,  1910  ...             ...  ...  ...  180 

Genus.     Paraplasma,  Seidelin,  1911                ...  ...  •••  180 

Sub-class.     Neosporidia,  Schaudinn             ...             ...             ...  ...  ...  181 

Order.     Myxosporidia,  Biitschli                 ...              ...              ...  ...  ...  181 

Order.     Microsporidia,  Balbiani                ...             ...             ...  ...  ...  184 

Order.     Actinomyxidia,  Stole.  ...             ...             ...             ...  ...  ...  187 

Order.     Sarcosporidia,  Balbiani                 ...              ...              ...  ...              ...  187 

Sarcospoiidia  oh?,et\tA  xn  Moxi           ...               ..             ...  193 

Order.     Haplosporidia,  C2i\x\\txy  zxxd  Me^mXy  i2><)()                ...  ...  ...  194 

Rhinosporidium  kinealyi,  Minchin  and  Fantham,  1905         ...  195 

Class  IV.     Infusoria,  Ledermiiller,  1763     ...             ...             ...  ...             ...  198 

Genus,     Balantidium,  Claparede  et  Lachmann  ...             ...  200 

Balantidium  coli,  Malmsten,  1857             ...  ...              ...  200 

Balantidium  minutum,  Schaudinn,  1899  ...              ...  204 

Genup.     Nyctotherus,  Leidy,  1849    •••             •••               ••             •••  204 

Nyclotherus faba,  Schaudinn,  1899            ...  ...             ...  205 

Nyctotherus giganieus,  P.  Krause,  1906    ...  ...              ...  205 

\Nyctotherus\africanus,QzsX&\\2.x\\,\iyd^  ...  ...             ...  206 

The  Chlamydozoa  ...            ...            ...            ...            ...  ...            ...  207 

Protozoa  Incert^  Sedis       ...            ...            ...            ...  ...            ...  210 

Sergenfella  ^ominis,  Brumpt,  igio            ...  ...             ...  210 


B.  Platyhelminthes  (or  Flat  Worms)  ...  ...  ...  ...  ...  211 

Classification  of  the  Platyhelminthes. 

Class  I.     Turbellaria  (or  Eddy  Worms)  ...  ...  ...  ...  212 

Order  I.     Rhabdoccelida      ...  ...  ...  ...  ...  ...  212 

2.  Tticladida  ...  ...  ...  ...  ...  ...  212 

3.  Polycladida         ...  ...  ...  ...  ...  ...  212 

Class    II.     Trematoda  (Sucking  Worms)  ...  ..  ...  ...  212 

III.     Cestoda  (Tapeworms)  ...  ...  ...  ...  ...  212 

Class  II.     Trematoda,  Rud.  ...  ...  ...  ...  ...  ...  212 

Development  of  the  Trematodes   ...  ...  ...  ...  .  .  222 

Biology       ...  ...  ...  ...  ...  ...  ...  ...  229 


CONTENTS  IX 

The  Animal  Parasites  of  Man — contd. 

Platyhelminthes— <r^«/^.  page 

Classification  of  the  Trematodes  of  Man. 

Order.     Z^z^i^w^a,  v.  Beneden,  1858          ...              ...              ...  ...  ...  230 

Sub-order.     Prostomata,  Odhner,  1905     ...              ...              ...  ...  ...  230 

Group.     Amphistoinata,  Rudolphi,  1801,  ep.,  Nitzsch,  1819        ...  ...  230 

Family.     FaramphistomidcE,  Fischoeder,  1901        ...  ...  ...  231 

Sub-family.     Paraviphislomince,  Fischoeder,  1 901  ...  ...  231 

Cladorchtince,  Fischoeder,  1901  ...  ...  ...  231 

Family.       G'ai/r^fl!'^V^Vfl'i^,  Stiles  and  Goldberger,  19 10  ...  ...  231 

Group.     Distomata,  Retzius,  1782    ...             ...             ...  ...  ...  231 

Family.     Fasciolidce,  Railliet,  1895            ...              ...  ...  ...  231 

Sub-family.     Fasciolince^  Odhner,  1910            ...  ...  ...  231 

Fasciolop since,  Odhner,  1910       ...  ...  ...  231 

Family.     Opisthorchiida,  Braun,  1901,  emend,  auctor.  ...  ...  232 

Sub-family.     Opisthorchiince,  Looss,  1899,  emend,  auctor.  ...  232 

MetorchmtcB,  Llihe,  1909              ...  ...  ...  232 

Family.     DicrocoeliidcB,  Odhner,  1910      ...              ...  ...  ...  232 

Heteroihyiidce,  Odhner,  1914    ...              ...  ...  ...  232 

Troglolreniid</:,  Odhner,  1914    ...              ...  ...  ••■  232 

Echmoslomidce^  Looss,  1902      ...              ...  ...  ...  233 

Sub-family.     Echitiostomince,  Looss,  1899       •••  ■••  •••  ^33 

HimasthliniE,  Odhner,  1910        ...  ...  ...  233 

Family.     Schistosomidce,  Looss,  1899        ...              ...  ...  ...  233 

The  Trematodes  observed  in  Man  ...            ...            ...  ...  ...  234 

Family.     FaramphistomidcB,  Stiles  and  Goldberger,  emend.  1910       ...  234 

Sub-family.     Cladorchiiuce,  Fischoeder,  1901           ...  ...  ...  234 

Genus,      ^f^a/j^^wmx.  Stiles  and  Goldberger,  1910  ...  ...  234 

^<7/5^^«/«.r  7fa/j-(7«/,  Stiles  and  Goldberger,  1910     ...  ...  234 

Family.      GastrodisciidcE               ...              ...              ...  ...  ...  236 

Genus.       Gasirodisais,  Lkt.,  1877     ...             ...  ...  ...  236 

Gastrodiscus  homiiiis,  'LeWi?,  dindiM.cConviQ\\f  \?>t6...  ...  236 

Family.     Fasciolidce,  Railliet,  1895           •••             •••  •••  ••  ^37 

Sub-family.     FasciolincB,  Odhner,  1910    ...             ...  ...  ...  237 

*     Genus.     Fasciola,  L.,  1758...             ...             ...  ...  ...  237 

Fasciola  hepatica,\u.,  Vji^Z          ...             ...  ...  ...  237 

Halzoun...             ...             ...             ...  ...  ...  242 

Fasciola giganti'ca,  Cobbold,  1856              ...  ...  ...  244 

Sub-family.     Fasciolopsitice,  Odhner,  1910               ...  ...  ...  245 

Genus.     Fasciolopsis,  Looss,  1898      ...             ...  ...  ...  245 

Fasciolopsis  buski,  Lank.,  1857  ...              ...  ...  ...  245 

Fasciolopsis  rathouisi,  Ward,  1903              ...  ...  ...  246 

Fasciolopsis goddardi.  Ward,  1910              ...  ...  ..  247 

Fasciolopsis fiilleborni,  Rodenwaldt,  1909  ...  ..  247 

^    Family.      Troglotremidce,  Odhner,  1914   ...              ...  ...  ...  249 

Genus.     Paragoninius,  BxeiMn,  1899...              ...  ...  ...  249 

Faragonitnus  ringeri^  Cobb.,  1880             ...  ...  ...  249 

Family.     Opisihorchiidcs,  Braun,  1 90 1      ...              ...  ...  ...  252 

Sub-family.     Opislhorchiince,  Looss,  1899                ...  ...  ...  252 

Genus.     Opisthorchis,  R.  Blanch.,  1845           •••  •••  "•  ^52 

Opisthorchis felineus,  Riv.,  1885                ...  ...  ...  252 

Genus.     Paropistkorchis,  Stephens,  1912         ...  ...  ...  255 

Paropisthorchis  caninus,  ^2ixV.Gry  1^12       ...  ...  ...  255 

Genus.     Amphimerus,  Barker,  191 2  (?)            ...  ...  •••  257 

Amphimerus  fiovejra,  Barker,  1912  (?)      ...  ...  ••■  258 

Genus.     Clonorchis,  Looss,  1907       ...             ...  ...  ...  258 


X  THE  ANIxMAL   PARASITES   OF   MAN 

The  Animal  Parasites  of  Man — contd. 
Platyhelminthes— r<?«i'a'. 

Clonorchis  sinensis,  Cobbold,  1875 
Clonorchis  endemicjis,^dit\Zy  1 883 
Sub-family.     MetorchiincB,  Liihe,  1 909     ...  ...  ... 

Genus.     Metorchis,  Looss,  1899,  emend,  auctor. 
Metorchis  truncatus,  Rud.,  1 81 9 
Family.     Hetei-ophyiidce,  Odhner,  1914... 
Genus.     Heterophyes,  Cobbold,  1866 

Heterophyes  heterophyes,  v.  Sieb.,  1 852 
Metagommtis,  Katsurada,  1913  ;  Yokogawa,  Leiper,  1913 
Melaoonimus  yokogawai,  Katsurada,  19 1 3 
Family.     Dicrocceliida,  Odhner,  1910 
Genus.     Dicroccclium^  Dujardin 

Dicroccelnim  dendrilicum,  Rud.,  18 19 
Family.     Eckinostomidis,  Looss,  1902 
Sub-family.     Echinostomince,  Looss,  1899 

Genus.     Echitiostoma,  Rud.,  1809;  Dietz,  1910 
Echinostoma  ilocanum,  G2iXt\^on,  lf)0?> 
Echinostoma  malayantirn,  Leiper,  1911     ... 
Sub-family.     Himasthlince,  Odhner,  1910 

Genus.     Artyfechinostornutn,  Clayton-Lane,  1915 

Artyfechinostomutn  siifrartyfex,  Clayton-Lane,  191 5 
Family.      Schistosomidce,  Looss,  1899       ...  ... 

Genus.     Schistosoma^  Weinl..  1858    ... 

Schistosoma  hivmatobitwi^  Bilharz,  1852     ... 
Schistosoma  mansoni,  Sambon,  1907 
Schistosoma  ;aponicu?f?,\s.2i\.s\M2idia,  1904... 

Class  IIL     Cestoda,  Rud.,  1808    ... 
Anatomy  OF  THE  Cestoda      ... 
Development  OF  THE  Tapeworms 
Biology 


Classification  of  the  Cestoda  of  Man. 

Order.     Pseudophyllidea,  Carus,  1863 

Family.     Dibothriocephalida,  Liihe,  1902 

Sub-family.     DibothriocephalincB,  Liihe,  1899 
Order.      Cyclophyllidea,  v.  Beneden 

Family.     DipylidiidcB,  Liihe,  1910    ... 

Hymenolepididce,  Railliet  and  Henry,  1909    ... 
Davaineidi?,  Fuhrmann,  1907 
Sub-family.     Davaineincey  Braun,  1900   ... 
Family.      Tc^niidce,  Ludwig,  1886     ... 

The  Cestodes  OF  Man 

Famiy.     Dibothriocephalidce 
Sub-family.     Dibothriocephalince 

Genus.     Dibothriocephalus,  Liihe,  1899 
Dibothriocephalus  latus,  L.,  1748 
Dibothriocephalus  cordatus,  R.  Lkt,,  1863 
Dibothriocephalus  parvus,  Stephens,  1908 
Genus.     Diplo^onoporus,  Lonnbrjj.,  1892 

Diplogonoporus  grandis,  R.  Blanch.,  1894 
Sparganuni,  Diesing,  1854 
Sparganum  mansoni,  Cobb.,  1883 
Sparganum  proliferum,  Ijima,  1905 


CONTENTS 


XI 


The  Animal  Parasites  of  yiAn—contd. 
Platyhelminthes — confd. 

Family.     Dipylidiidce,  Liihe,  1 910 

Genus.     Dipylidium,  R.  Lkt.,  1863... 
Dipylidium  canhuun,  L.  1 758    ... 
Family.     ^;/wg«^/^/za?'/fl('^,  Railliet  and  Henry,  1909 
Genus.     Hymenolepis,  Weinland,  1858 
Hymenolepis  nana,  v.  Sieb.,  1852 
Hytnenokpis  diminuta,  Rud.,  18 19 
Hymenolepis  lanceolata,  Bloch,  1782 
Family.     Davaineidce,  Fuhrmann,  1907   ... 
Sub-family.     Davaineina;,  Braun,  1900    ... 
Genus.     Davainea,  R.  Blanch.,  189 1 

Davainea  madagascariensis,  Davaine,  1869 
Davainea  (?)  asiaiica,  v.  Linst.,  1901 
Family.      Tceniid(C,  Ludwig,  1886 
Genus.      Tanm,  L.,  1758    ... 

Tania  solniin^'L,.,  p.  p.^  I'jd'j     ... 
Cysticercus  acanthotrias,  Weinland,  1858... 
Tli'wza  ^r^///««'z,  Stephens,  1908 
Tcenia  marginata,  Batsch.,  1786 
Tienia  serrata,  Goeze,  1782 
Tcenia  crassicollis,  Rud.,  1810    ... 
Tc^nta  saginala,  GoezQ,  i'j2>2 
Ticnia  africana,  \.  Linst.,  1900... 
Tcenia  conftisa,  Ward,  1896 
Tcenia  echinococcHs,  v.  Sieb.,  1853 

Structure  and  Development  of  Ecchinococcus  (Hydatid) 
Echinococctts  mtiltilocularis  (Alveolar  Colloid) 
Serum  Diagnosis  of  Echinococcus     ... 

C.  Nemathelmintljes 

Class.     Nematoda 

Anatomy  of  the  Nematodes 
Development  of  the  Nematodes 


Classification  of  the  Nematoda. 
Family.    Anguillulidic,  Gervais  and  van  Beneden,  1859 
Angiostomidcd,  Braun,  1895 
Gnathostoviidce 
Dractmculidce,  Leiper,  19 12 
Filar iidce,  Claus,  1885 
TrichinellidLe,  Stiles  and  Crane,  1 9 10 
Dioctophyinidcc 
Strongylida,  Cobbold,  1864 
Physalopteridce 
Ascaridce,  Cobbold,  1864 
Oxyuridce 

The  Nematodes  observed  in  Man  ... 
Family.     Anguillulidcc 

Genus.     Rhabdiiis,  Dujardin,  1845  ••• 
Rhabditis  pellio,  Schneider,  1866 
Rhabditis  niellyi,  Blanchard,  1885 
Rhabditis,  sp. 
Genus.     Anguillula,  Ehrenberg,  1826 
Anguillula  aceli,  Miiller,  1783  ... 


xu 


THE   ANIMAL   PARASITES   OF   MAN 


The  Animal  Parasites  of  yiA^—contd. 
Nemathelminthes— <r^w/df. 

Genus.     Anguillulmay  Gervais  and  Beneden,  1859 
Anguillulina piitrefaciens,  Kiihn,  1879     ... 
Family.     Angioslomida^  Braun,  189'; 
Genus.     Strongyloides,  Grassi,  1879 

Strongyloides  stercoralii,  Bavay,  1877 
Family.     GnathostomidiZ 

Genus.      Gnathostotna,  Owen,  1836  ... 

Gnathostoma  siamense,  Levinson,  1889    ... 
Gnathostoma  spinigerum,  Owen,  1836 
Family.     Dracuncjilidie,  Leiper,  191 2 

Genus.     Draciincuhis^  Kniphoff,  1759 

Dracunadus  jjudinensis,  Velsch,  1674 
Genus  (of  Crustacea).     Cyclops^  Muller,  1776... 
Family.     Filariida 
Sub-family.     Filariincc 

Genus.     Filaria,  O.  Fr.  Muller,  1787 
Filarta  bancrofti^  Cobbold,  1877 
Filar ia  deviarquayiy  Manson,  1895 
Filarta  taniguchi,  Penel,  1905  ... 
Filaria  (?)  conjimctivce,  Addario,  1 885     ... 
Group.     Agavio filaria.  Stiles,  1906  ... 
Agamo filaria  georgiana 
Agafjiofilaria  palpebralis.  Pace,  1 867  («^^  Wilson,  1844) 
Aganiofilaria  oculi  humani,  v.  Nordmann,  1832 
Agavwfilaria  labialis,  Pane,  1864 
Filaria  (?)  romanormn-orientalis ,  Sarcani,  1888 
Filaria  (?)  kilimarce,  Kolb,  1 898 
Filaria  (?)  sp.  ? 
Genus.     Setaria,  Viborg,  1795 

Setaria  eqtiina,  AhWAg.,  1789     ... 
Genus.     Loa,  Stiles,  1905  ... 
Loa  loa,  Guyot,  1778    ... 
Genus.     Acanthocheilonema,  Cobbold,  1870    ... 
Acanthocheilonema  Persians,  Manson,  1 891 
Genus.     Dirqfilaria,  Railliet  and  Henry,  1911 
Diro filaria  magalliaesi,  R.  Blanchard,  1895 
Sub-family.     Onchocercince,  Leiper,  19 ii 
Genus.     Onchocerca,  Diesing,  1841   ... 

Onchocerca  volvulus,  R.  Leuckart,  1 893  ... 
Family.  Trichinellidce,  Stiles  and  Crane,  1910  ... 
Sub-family.      Trichurince,  Ransom,  191 1 

Genus.     Trichuris,  Roderer  and  Wagler,  1761 
Trichuris  trichiura,  Linnreus,  1 761 
Sub-family.      TrichinellincE,  Ransom,  1911 
Genus.      Trichinella,  Railliet,  1895  ••* 
Trichinella  spiralis,  Owen,  1835 

History  of  the  Development  of  Trichinella  spiralis 
Family.     Dioctophymidct 

Genus.     Dioctophyme,  Collet-Megret,  1802     ... 
Dioctophyme  gigas,  Rudolphi,  1802 
Family.     Strongylidce  ... 
Sub-family.     Metastrongylince,  Leiper,  1908 
Genus.     Metastrongylus,  Molin,  1861 
Melastrongylus  apri,  Gmelin,  1789 
Sub-family.      Trichosirongylince,  Leiper,  1908 


CONTENTS  XUl 

The  Animal  Parasites  of  Man — contd. 

Nemathelminthes — contd.  pagr 

Genus.      Trichostrongylus,  Looss,  1905            ...  ...  ...  434 

Trichostrongyhis  instabilis,  Railliet,  1893  •••  •••  434 

Jrichostrongylus probohirus^  Railliet,  1896  ...  ...  435 

Ti'ichosirongylus  vilriinis,  Looss,  1905     ,..  ...  ...  435 

Genus.     Hcemoiichtis,  Cobb.,  1898    ...              ...  ...  ...  436 

Hcemonchus  conloi'tus,  Rudolphi,  1803  ;  Cobb.,  1898  ...  436 

Genus.     Nematodirus^  Ransom,  1907,  emend.  Railliet,  1912      ...  438 

Sub-genus.     Mecisiorirrus,  Railliet,  1912        ...  ...  ...  438 

Mecis/oci}'rtisfordi,T>3imt\s,i^o2>              ...  ...  ...  438 

Sub-family.     Ancylostoinimc,  Raillet,  1909I              ...  ...  ...  438 

Group.      CEsophagostomecc,  Railliet  and  Henry,  1909     ...  ...  ...  439 

Genus.      Ternidens,  Railliet,  1909     ...              ...  ...  ...  439 

Ternidens  deminutus,  Railliet  and  Henry,  1905  ...  ...  440 

Genus.      (Esophagostofiiuni,  Molin,  1861           ...  ...  ...  441 

CEsophagostoinuiii  bniniptiy  Railliet  and  Henry,  190';  ...  441 
CEsophagostofiiHtn  stephanostonmm  var.  thomasi,  Railliet  and 

Henry,  1909       ...              ...              ...  ...  ...  443 

(Esophagostovium  apiostonnini,  Willach,  1891  ...  ...  444 

Group.     AncylostomecE,  Railliet  and  Henry,  1909           ...  ...  ...  445 

Genus.     Ancylostoma,  Dubini,  1843,  emend.  Looss,  1905  ...  445 

Ancylosioma  duodenale,T)\x\nn\,  1843        ...  ...  ...  445 

Ancy loslovi a  ceylanicuvi,  \aOQ%%,  19 ii         ...  ...  ...  456 

Ancylostotna  brazilietise^Qoxn&z  ^^Y^ixxz,  \<^\o  ...  ...  456 

Group.     .5/^«i7i'/tfz//<?cr,  Railliet  and  Henry,  1909             ...  ...  ...  456 

Genus.     Necator^  Stiles,  1903              ...             ...  ...  ...  457 

Necator  aniericamis,'$i\\\^%^  if^02                 ...  ...  ...  457 

Necator  exilidens,  Cummins,  1912             ...  ...  ...  459 

Ancylostomiasis     ...              ...              ...  ...  ...  459 

Group.     6)/ «^a/W(?^,  Railliet  and  Henry,  1909                ...  ...  ...  459 

Genus.     Syngamus,  von  Siebold,  1836             ...  ...  ...  459 

Sytigamus  kingi,  Leiper,  1913...             ...  ...  ...  459 

Family.     Physalopteridcc              ...              ...              ...  ...  ...  460 

Genus.     Physaloptera^  Rudolphi,  18 19             ...  ...  ..  460 

Physaloptera  caucasica,  \.  Uiwhto^,  1^02  ...  ...  ...  461 

Physaloptera  mordens,  Leiper,  1907            ...  ...  ...  461 

Family.     Ascaridce,  Cobbold,  1864            ...              ...  ...  ...  461 

Sub-family.     Ascarince                 ...              ...              ...  ...  ...  461 

Genus.     Ascaris,  L,.,  1758...             ...             ...  ...  ...  461 

Ascaris /umbricoi'des,  Li. ,  ly S^    •••             •••  •••  •••  4^3 

Ascaris  sp.      ...              ...              ...              ...  ...  ...  465 

Ascaris  iexana,  Sm\\.h  Qt  Goeih,  igi/^       ...  ...  ...  465 

Ascaris  martitma,  Leuckart,  1876              ...  ...  ...  465 

Genus.      Toxascarzs,  Leiper,  1907     ...             ...  ...  ...  465 

Toxascaris  h'/uba/a,  Railliet  and  Henry,  191 1  ...  ...  466 

Genus.     Be/ascaris,  Leiper,  1907      ...             ...  ...  ...  466 

Belascaris  cati,  SchxdiVtk,  1788     ...              ...  ...  ...  466 

Belascaris  tnarginata,  Rudolphi,  1802       ...  ...  ...  466 

Genus.     Lagocheilascaris,  Leiper,  1909            ...  ...  ...  466 

Lagocheilascaris  minor.,  Leiper,  1909        ...  ...  ...  467 

Family.     Oxyurid(V      ...             ...             ...             ...  ...  ...  467 

Genus.     Oxytiris^  Rudolphi,  1803     ...              ..,  ...  ...  467 

Oxyuris  vermiciilaris,  Linnceus,  1767        •••  •••  •••  4^7 

Family.     Merniiihid<€ ...              ...              ...              ...  ...  ...  469 

Genus.     Merniis,  Dujardin,  1845       ...              ...  ...  ...  469 

Mermis  Aominis  oris,  L,eidy,  18^0             ...  ...  ...  469 


xiv  THE  ANIMAL   PARASITES   OF   MAN 

The  Animal  Parasites  of  Man — contd, 

Nemathelminthes — contd.  page 

Agatnomermis^  Stiles,  1903          ...  ...  ...  •••  47° 

Agamomermis  restiforniis,  Leidy,  1 880  ...  ...  ...  47° 

Technique  ...            ...            ...            ...            ...  •••  •••  •••  47i 

D.  Acanthocephala,  Rud....            ...            ...            ...  ...  ...  ••  475 

Echinoj-hyrtchifs  gigas,  Goeze,  1782  ...  ...  ...  477 

Echiuorhyitchus  hominiSy'LaxohX,  \2>$g  ...  ...  ...  47^ 

Echinorhynchus  moniliformis,  V>x&xii%^x,  \^\.<^  ...  ...  478 

E.  Gordiidae        ...            ...            ...            ...             •.  ••  •••  •••  479 


F.  Hirudinea  s.  Discophora  (Leech)             ...            ...  ...  ...  ...  480 

Family.     6';/aMo^fl'^///^^  (Leeches  with  Jaws)  ...  ...  ...  481 

Genus.     Hirudo,  L.,  1758  ...             ...  ...  ...  ...  481 

Hirudo  medicinalis,L..y  lT^2i      ...  ...  ...  ...  481 

Hirudo  troctina,  ]o)M\%\.or\,  1816  ...  ...  ...  482 

Genus.     Limnatis,  Moq.-Tandon,  1826  ...  ...  ...  482 

Limnatis  nilolica,  Savigny,  1820  ...  ...  ...  482 

Genus.     Hicmadipsa,  Tennent,  1 86 1...  ...  ...  ...  482 

Family.     Khynclwbdellidce  (Leeches  with  Rostrum)  ...  ...  482 

Genus.     Htetnentaria,  de  Filippi,  1849  ...  ...  ...  482 

Hamentaria  officinalis y  dQ¥\\\^^\  ...  ...  ...  482 

Genus.     Placobdella,  R.  Blanchard    ...  ...  ...  ...  482 

Placobdella  calenigera,  M.o(\.-l!2indoTi  ..  ...  ...  482 


G.  Arthropoda  (Jointed-limbed  Animals)       ...  ...  ...  ...  ...  483 

^.  Arachnoidea  (Spiders,  Mites,  etc.)  ...  ...  ...  ...  ...  483 

Order.     Acarina  (Mites)  ...  ...  ...  ...  ...  ...  484 

Family.      Trombidiidcc  {K\xx\xi\x\^M\\.^%)   ...  ...  ...  ...  485 

Genus.      Ti ombidiian,  'L.dXx^'iWe.  {^cxidi  Lepttis)  ...  ...  ...  485 

Leplus  atitumnalis,  ShsiViy  1790...  ...  ...  ...  485 

Trombidium  tlalsahuate^  Lemaire,  1867   ...  ...  ...  486 

Akamiishi  ox  Kedani    ...  ...  ...  ...  ...  487 

Family.      7"<j^ra«j/r/^?V/c?  (Spinning  Mites)...  ...  ...  ...  488 

Genus.     Tecrauychtts,  Dufour  ...  ...  ...  ...  488 

Telranyc/ius  moleslissimits,  Weyenhexgh,  1886        ...  ...  488 

Tetranychus  ielarius,  L.,  1758,  var.  lusseoins,  Koch  ...  488 

Family.      Tarsonemidcc  ...  ...  ...  ...  ...  488 

Genus.     Pediculoides  ...  ...  ...  ...  ...  489 

Fediculoides  ventricosus,^eviY>oxi,  i%^o   ...  ...  ...  489 

Genus.     Neph7-ophages         ...  ...  ...  ..  ...  490 

Nephrophages  sanguinariusy  Miyake  and  Scriba,  1893  ...  490 

Family.     Eupodidce      ...  ...  ...  ...  ...  ..  491 

Genus.      Tydeus,  Koch        ...  ...  ...  ...  ...  491 

7 ydeus  mo/eslus,  Mon\ez,  iSSg  ...  ...  ...  ...  491 

Family.     Gainasidcc  (Coleopterous  or  Insect  Mites)  ...  ...  491 

Genus.     Dermanyssus,  Duges  ...  ...  ...  ...  492 

Derjnatiyssus  gallince,  di^  Gqqx,  1778         ...  ...  ..  492 

Dertnanyssns  hirundinis,  Hermann,  1804  ,,.  ...  492 

Genus.     Holothyrus  ...  ...  ...  ...  ...  493 

Holothyrus  coccinella,  Gervais,  1842         ...  ...  ...  493 


CONTENTS 


XV 


The  Animal  Parasites  of  Man — contd. 

Arthropoda-  <r^«/«'. 

Family.     Ixodidce  (Ticks) 
Classification  of  Zr(?(a?iVfe 
Synopsis  of  Genera 

Genus.     Ixodes,  Latreille    ... 

Ixodes  redtivmsj  L.,  1758 

Ixodes  holocyc his,  'ii&xxrcia.nn,  1899 

Ixodes  hexagonus,  Leach,  1815... 
Genus.     Amhlyomma,  Koch 

Amblyomma  coyennejise,  Koch,  1 844 

Amblyotmna  aniericana,  Linnseus 

Amblyomma  maculatzim,  Koch  ... 
Genus.     Hyalomma,  Koch 

Hyalom  m  a  cegyptiu  /// ,  L. ,  1 7  5  8  , . . 
Genus.     Htzmaphysalis,  Koch 

Hcemaphysalis  punctata,  Canestrini  and  Fanzago,  1877-1878 
Genus.     Dermacentor,  Koch 

Dermacenior  retictdatus,  Fabricius,  1 794... 

Dermacentor  venuslus,^a.V)\is,     ... 

Dermacentor  occtdenta/is,  'NeumsLDn 

Dermacentor  variabilis,  Say 
Genus.     Margaroptis^  Karsch 

Margaropus  anmdatus  australisy  Fuller  ... 

Margaropus  juicroplus,  Canestrini 
Genus.     Rhipicephalus^  Koch 

Rhipicephalus  sanguineus^  Latreille,  1804 
Neumann's  Table  of  Species  of  ^r^ai- 
Genus.     Argas,  Latreille    ... 

Argas  rejlexus,  Fahiicius,  iyg4  ... 

Argas  persicus,  Fischer  de'Wa.ldheim^  1824 

Argas  brumpti,  Neumann 

Argas  chine  he.,  Gervais,  1844 
Genus.      Ornithodorus,  Koch 

Ornithodorus  7noubata,  Murray,  1877 

Ornithodorus  savignyi,  Audouin,  1827 

Ornithodorus  coriaceus,  Koch 

Ornithodorus  talaje,  Guerin,  1849 

Ornithodorus  turicata,  Duges,  1876 

Ornithodorus  tholozani,  Laboulbene  and  Megnin,  1882 

Ornithoaorus  megnini,  Duges,  1883 
Family.     Tyroglyphida 
Sub-family.      Tyroglyphince 

Genus.     Aleurobius,  Canestrini 

Aleurobius  {lyroglyphus)  farince,  de  Geer  (part),  Koch 
Genus.     Tyroglyphus,  Latreille 

Tyroglyphus  siro,  L.,  1756 

Tyroglyphus  longior,  Gervais,  1 844 

Tyroglyphus  minor  var.  castellani.  Hirst  ... 
Genus.     Glyciphagus,  Hering,  1838... 

Glyciphagus  prunorum^  Hering,  and   G.  domesticus,  de  Geer 

Glyciphagus  cursor,  Gtivaxs 

Glyciphagus  buski,  Murray 
Genus.     Rhizoglyphus,  Claparede,  1869 

Rhizoglyphus  parasiticus ,  Dalgetty,  1901... 


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513 
513 
513 

514 


XVI 


THE  ANIMAL  PARASITES  OF  MAN 


The  Animal  Parasites  of  Man — contd. 
Arthropoda— fc;//^. 

Genus.     Histiogaster,  Berlese,  1883  ... 

Histiogaster  {^entomophagus  ?)  spermaticus,  Trouessart,   1 900 
Genus.     Cheyletus 

Cheyletus  mericourti.  Lab. 
Family.     Sarcoptidcc  (Itch  Mites) 
Sub-family.     Sarcoptince 

Genus.     Sarcoptes,  Latreille 

Sarcoptes  scabiei,  de  Geer,  1778 
Sarcoptes  minor,  PHirstenberg,  1861 
Family.     Demodicidce  (Mites  of  the  Hair-follicles)... 
Genus.     Deiriodex,  Owen    ... 

Demodex  follictilortim,  Simon,  1842 
Order.     Pentastomida  ... 

Family.     Linguatulidts 

Genus.     Ltt2guaiula,  Frohlich 

Linguatula  rhinaria,  Pilger,  1 802 
Genus.     Forocephahts 

Porocephalus  constrictiis,  v.  Siebold,  1852 
B.   Insecta  (Hexapoda)     ... 

Classification  of  the  Hexapoda. 

(i)  Aptera 

(2)  Neuroptera 

(3)  Orthoptera 

(4)  Thysanoptera 

(5)  Hemiptera 

(6)  Diptera    ... 

(7)  Lepidoptera 

(8)  Hymenoptera 

(9)  Coleoptera 
Order.     Rhyncota 

(a)  Rhyncota  aptera  parasitica 
Family.     Pediculidcc  (Lice) 

Genus.     Pedictilics,  Linnaeus 

Pediculus  capitis,  de  Geer,  1778 
Pediculns  vestimenti,  Nitzsch,  1818 
Genus.     Phthiritis,  Leach  ... 

Phthirius  itiguinalis,  Redi,  1668 
{b)  Rhyncota  hemiptera 

Family.     Acanthiadce  ... 

Genus.      Cimex,  Linnseus    ... 
Cimex /ectu/arius,  hmriceus 
Cirnex  ro/undatus,  Signoret,  iHs2 
Cifnex  colu7/ibarius,  ]enyx\s 
Cimex  ciliatus,  Eversmann,  1841 
Family.     Reduviidcr    ... 

Genus.     Conorhimis,  Lap.... 

Conor  hi  Hus  megisius,  Burrn. 

Conof  hinus  sanguisuga,  Lee.  (Blood-sucking  Cone-nose) 

Conorhimis,  sp.  novum  (Monster  Bug) 

Conorhimis  rubrofasciatus,  de  Geer  (Malay  Bug)    ... 

Conorhimis    renggeri,    Herr-Schaff    (Great    Black     Bug 

Pampas) 
Conor hinus  variegatus  (Variegated  Cone-nose) 


CONTENTS  XVU 

The  Animal  Parasites  of  M\^—contd. 

Arthropoda— tw//<^.  page 

Conorhinus  nigrovarius               ...  ...  ...  ...  539 

Cotiorhinus  protractus ...              ....  ...  ...  ...  539 

Genus.     Keduvius,  etc.       ...             ...  ...  ...  ...  539 

Reduvius personatus,  Linne         ...  ...  ...  ...  539 

Coriscus  subcoleoptratuSyYsAxhy,  iZyj  ...  ...  ...  540 

Rasahus  bigntlatuSySa-y,  i^'^i    ...  ...  ...  ...  540 

Melanolestes  viorio,  Erichson,  1848  (non- Walker)  ...  ...  540 

Melanolesies  abJoniinalis,  Htxx'ichSch'i.nQX,  iZ^%    ...  ...  540 

Phonergates  b  i  color  i pes ...             ...  ...  ...  ...  541 

Family.     Aradida        ...             ...             ...  ...  ...  ...  541 

'                                                    Dysodius  lunatus,  Fabr.  (Pito  Bug)  ...  ...  ...  541 

The  Ochindundu  ...             ...  ..:  ...  ...  541 

Family.     Lygczidce        ...              ...              ...  ...  ...  ...  541 

Lyctocoris  cavipesiris,Y2\yc\c\\x%  ...  ...  ...  ...  541 

Rhodinus prolixusi  Slal,  1859   ...  ...  ...  ...  541 

Order.     Orthoptera      ...             ...             ...             ...  ...  ...  ...  542 

Locusts  Injuiious  to  Man    ...             ...             ...  ...  ...  ...  542 

Order.     Coleoptera       ...             ...             ...             ...  ...  ...  ...  542 

Silvanus surinamensis,  Linnceus  (Saw-toolhed  Grain  Beetle)...  542 

Order.     Diptera           ...             ...             ...             ...  ...  ...  ..  543 

Ap/iampiera  ox  Siphonap/era  (Fleas)                 ....  ...  ...  ...  543 

Family.     Sarcopsy//id(v  Qigi^exs)                ...  ...  ...  .  ...  543 

Genus.     Dermatophilus^  Guerin        ...  ...  ...  ...  544 

Dermalophilus  cceca/a,Y.i\dex\.  ...  ...  ...  ...  544 

Dermatophihts penetrans,  L.,  1 758  (Jigger,  Chigoe)  ...  544 

Genus.      Echidnophaga,  Olliflf           ...  ...  ...  ...  544 

Echidnophaga  gallinacea,  Westwood  (Chigoe  of  Fowls)        ...  544 

Family.     PuHcidce  (True  Fleas)...             ...  ...  ...  ...  545 

Genus.     Pulex,  Linn.         ...             ,..  ...  ...  ...  545 

Pulex  irritans,  L.,  1758              ...  ...  ...  ...  545 

Genus.     Xenopsylla,  Glink...             ...  ...  ...  ...  546 

Xenopsylla  cheopis,^o\.\i%f^\\A    ...  ...  ...  ...  546 

Xenopsylla  brastliensis,  Baker    ...  ...  ...  ,,,  §47 

Genus.     Ctenocephalus,  Kolenati       ...  ...  ...  ...  547 

Genus.     Hoplopsyllus,  Baker              ..,  ...  ..,  ...  547 

Hoplopsyllus  atiofnalns,  Baker    ...  ...  ...  ...  caj 

Genus.     Ceratophyllus,  Cen\\s           ....  ...  ...  .,,  547 

Ceratophyllus fascialusy  Bosc      ...  ...  ...  ...  547 

Genus.      Ctenopsylla,  Kolenati           ...  ...  ...  ...  r48 

Genus.     Hystrichopsylla,  Taschenberg  ...  ...  ...  ^48 

Pulex  pallipes                ...              ..,  ...  ...  ...  ^^g 

Systematic  Anatomical  and  Biological  Remarks  on  Mosquitoes    ...  548 

CuLiciD^  OR  Mosquitoes    ...            ...            ...  ...  ...  ..,  cee 

The  Classification  of  Culicida                  ...              ...  ...  ...  ...  c6i 

Notes  on  the  Different  Genera                ...             ...  ...  ...  . _  c66 

Sub-family.     Anophelina             ...              ...  ...  ...  ...  pgg 

Genus.     Anopheles,  Meigen                ...  ...  ...  ...  egg 

Genus.     Alyzomyia,  Blanchard  ;  Grassia,  Theobald     ...  ...  567 

Genus.     Neoinyzomyia.,  Theobald      ...  ...  ...  ...  567 

Genus.     Cycloleppteron,  Theobald    ...  ...  ...  ...  eg* 

Genus.     Feltinella,  Theobald             ...  ...  ...  ...  ^i^-j. 

Genus.     Stethojnyia,  Theobald          ...  ...  ...  ...  cfir 

Genus.     Pyretophortis,  Blanchard  ;  Hoiuardia,  Theobald  ...  567 

Genus.     Myzorhynchella,  Theobald  ...  ...  ...  ...  ^6&. 


XVlll 


THE  ANIMAL  PARASITES  OF  MAN 


The  Animal  Parasites  of  Man — contd. 
Arthropoda—  contd. 

Genus. 


Manguinhosia,  Cruz  (in  Peryassu)     ... 
Genus.     Chrystya,  Theobald  ..." 

Genus.     Lophoscelomyia,  Theobald  ... 
Genus.     Arribalzagia,  Theobald       ...  ...  ... 

Genus.     Myzorhyiichus^^\zxiQ}ci?i\^\  ^<?jjza,  Theobald... 
Genus.     Nyssorhynchus,'^\2S\c}[\2ixd>.\  Laverania^TYi^oh^X^ 
Genus.     Celliay  Theobald  ... 
Genus.     Neocellia,  Theobald 
Genus.     Kertesziuy  Theobald 
Genus.     Manguinhosia,  Ciuz 
Genus.     Chagasia,  Cruz 
Genus.     Calveriina,  Ludlow 
Genus.     Birbnella,  Theobald 
Sub-family.     Megarhinina 

Genus.     Alegar hinus,  Robineau  Desvoidy 
Genus.      Toxorliynchites,  Theobald  ... 
Sub-family.     Culicince ... 

Genus.     Mucidus^  Theobald 
Genus.     Psorophora^  Robineau  Desvoidy 
Genus.    Janthinosoma,  Arribalzaga  ... 
Genus.     Stegomyia^  Theobald 

Stegomyia  fascial  a,  Fabricius  (Yellow  Fever  Mosquito) 
Stegoviyia  scutellaris.  Walker 
Genus.      Theobaldia,  Neveu-Lemaire 
Theobaldinella^  Blanchard 
Theobaldia  antmlata,  M.G.\gQn 
Genus.     Culex,  Linnaeus 
Genus.     Melanoconion^  Theobald 
Genus.     Grabhamia,  Theobald 
Genus.        Psetidotceniorhynchus,      Theobald  ;        Tccniorhynchus^ 

Theobald,  non-Arribalzaga 
Genus.       Tceniorhynchus^    Arribalzaga ;     Mansonia,    Blanchard ; 

Panopliiesy  Theobald 
Genus.     Chrysoconops,  Goeldi 
Other  Nematocera 

Family.     Simulidce 
Family.     ChirononiidcE  (Midges) 
Sub-family.      CeratopogonincE 
Family.     Psychodidce  (Owl  Midges) 
Brachyera  (Flies) 

Family.     Phoridcs 

AphiochcEta  ferruginea,  Brun 
Phora  ruJipeSy  'M.^\g.     ... 

Family.     Sepsidcz 

Piophila  casei,\,. 
Family.     ^S^r/^zflfe  (Hover  and  Drone  Flies) 
Family.     Drosophilidce 

Drosophila  melanogastery  Br.      ...  ...  ... 

Family.     Muscidce 

Teichomyzafusca,  Macq. 

Homalomyia  canicularis^'Li.,  &{c. 

Honialomyia  scalaris^  Fabr. 

Anthomyta  desjardensii,  Macq.  ...  ...  ... 

Hydrotaa  meteor ica^  L. 
Cyrtonetira  siabulans    ... 


PAGE 

568 
568 
568 
568 
568 
569 
569 
569 
569 
569 
570 
570 
570 
570 
570 
S70 
571 
571 
571 
571 
571 
574 
575 
575 
575 
575 
575 
576 
576 

576 

577 
577 
577 
577 
579 
580 
581 
582 
582 
583 
583 
583 
583 
583 
584 
584 
584 
584 
584 
585 
585 
585 
585 


CONTENTS  XIX 

The  Animal  Parasites  of  Man — contd. 

Arthropoda—  contd.  page 

yl/wi-^ro!  ^/^w<?j//Vfl!,  Linn.  (Common  House-fly)  ...  ...  585 

Genus.     Chrysomyia,  Rob.  Desv.      ...  ...  ...  ...  587 

Chrysoniyia     {Compsomyid)     macellaria,      Fabr.  ;      Lucilia 

macellaria,  Fabr.       ...             ...  ...  ...  ...  587 

Chrysomyia  virtdula,'Roh.'De%v.  ...  ...  ...  588 

Genus.     Lucilia^  Rob.  Desv.             ...  ...  ...  ...  588 

Lucilia  nobiliSy  M&\g.  ...             ...  ...  ...  ...  588 

Genus.     /^<:«i7j(7wa,  Brauer  and  v.  Bergenstamm  ...  ...  588 

Genus.     Sarcophaga,  Mg.    ...             ...  ...  ...  ...  589 

Sarcophaga  carnaria,!^.^  iT^^    ...  ...  ...  ...  589 

Sarcophaga  magnifica^  Schiner,  1862  ...  ...  ...  589 

Sarcophaga  chrysotoma,V^\tdL     ...  ...  ...  ...  590 

Sarcophaga  p/inthopygaf  Wied    ...  ...  ...  ...  590 

Ochromyia  anthropophaga^    E.   Blanch.  ;     Cordylobia  arthro- 

phaga^  Griinberg               ...  ...  ...  ...  590 

Auchmeromyia{Bengalid)  depressa  i^z^tx)  ...  ...  591 

Genus.     Cordylobia^  Griinberg,  1903  ...  ...  ...  591 

Cordylobia griinbergiy  Donitz      ...  ...  ...  ...  591 

Cordylobia  anthropophaga^  Griinberg  ...  ...  ...  592 

Lund's  Larva                ...             ...  ...  ...  ...  593 

Auchmeromyia  luleolaj  FahiicivLS  ...  ...  ...  593 

Family.     Oestrida        ...             ...             ...  ...  ...  ...  594 

Cutaneous  Oestridce             ...             ...             ...  ...  ...  ...  595 

Genus.     Hypoderma.,  Latreille            ...  ...  ...  ...  595 

Hypoderi7ia  bonis,  de  Geer           ...  ...  ...  ...  595 

Hypoderrna  lineata,  di&V\\\tts     ...  ...  ...  ...  596 

ILypoderma  diana^  ^i2Mtt           ...  ...  ...  ...  596 

Genus.     Dermatobia,  Brauer              ...  ...  ...  ...  596 

Dermatobia  cyaniventris^  Macq ....  ...  ...  ...  596 

Cavicolous  Oestridce              ...             ...             ...  ...  ...  ...  598 

Genus.     Oestrus,  Linnaeus  ...             ...  ...  ...  ...  598 

Oestrus  {Cephalo?nyia)  ovis,  L.    ...  ...  ...  ...  598 

Gastricolous  Oestridce            ...             ...             ...  ...  ...  ...  599 

Genus.     Gastrophilus,  Leach              ...  ...  ...  ...  599 

Biting-mouthed  and  other  Noxious  Diptera  which  may  be  Disease  Carriers     600 

Family.      Tabanidcx  (Gad  Flies)                 ...  ...  ...  ...  600 

Family.     Asilidce  ( Wolf  Flies)    ...             ...  ...  ...  ...  602 

Family.     Leptidce          ...             ...             ...  ...  ...  ...  603 

Blood-sucking  Muscidce        ...             ...             ...  ...  ...  ...  603 

Genus.     G/ossina,  Westwood             ...  ...  ...  ...  603 

Glossina pa/palis,  Rob.  Desv.       ..  ...  ...  ...  607 

Glossina  morsitans,  Westwood    ...  ...              ..  ...  608 

Genus.     Stomoxys,  Geoffroy               ...  ...  ...  ...  609 

Genus.     Lyperosia,  Rondani               ...  ...  ...  ...  610 

Pupipara  ox  Eproboscidce      ...             ...             ...  ...  ...  ...  .611 

Insects  and  Epidemic  Poliomyelitis            •••.         •••  •••  •••  ...  612 

Addenda             ...            ...            ...            ...            ...  ...  ...  ...  613 

Akamushi  or  Kedani  Sickness    ...             ...             ...  ...  ...  ...  613 

Ticks. — African  Tick  Fever        ...             ...             ...  ...  ...  ...  613 

Tick  Paralysis                ...              ...              ...              ...  ...  ...  ...  613 

Diptera. — Psychodidce  ...              ...              ...              ...  ...  ...  ...  613 

PulicidcE. — Dermatophihis  {Sarcopsylla)  penetrans,  or  the  "Jigger"   ...  ...  613 

Brachycera. — Leptidce  ...             ...             ...             ...  ...  ...  ...  613 

Myiasis            ...             ...             ...             ...             ...  ...  ...  ••.  615 


XX  THE  ANIMAL  PARASITES  OF  MAN 

The  Animal  Parasites  of  Man — contd. 
Arihropodsi—con^d. 

Auricular  Myiasis  ...  ...  ...  ...  ...  ...  ••      615 

Body,  Head,  and  Clothes  Lice   ...  ...  ...  ...  ...  ...     615 


PAGl 


SUPPLEMENT:  CLINICAL  AND  THERAPEUTICAL  NOTES  617 

Protozoa             ...           ...           ...  ...  ...  ...  ...  ..  617 

Introduction           ...             ...             ...  ...  ...  ...  ...  ...  617 

I. — Amcebic  Dysentery          ...  ...  ...  ...  ...  ...  618 

II. — Trypanosomiases            ...  ...  ...  ...  ...  ...  620 

African  Sleeping  Sickness  ...  ...  ...  ...  ...  ...  620 

South  American  Trypanosomiasis  ...  ...  ...  ...  ...  623 

III. — Flagellate  Diarrhcea  and  Dysentery  ...  ...  ...  623 

IV.— Leishmaniases                 ...  ...  ...  ...  ...  ...  626 

A.  Kala-azar         ...             ...  ...  ...  ...  ...  ...  626 

Indian            ...             ...  ...  ...  ...  ...  ...  626 

Infantile         ...             ...  ...  ...  ...  627 

B.  OtientAl  SotCt  due  to  LeisAmama  tropica  ...  ...  ...  627 

Naso-oral  (Espundia)   ...  ...  ...  ...  ...  ...  628 

V. — Spiroch/ETOSes  ...            ...  ...  ...  ...  ...  ...  629 

A.  Relapsing  Fevers           ...  ...  ...  ...  ...  ...  629 

B.  Yaws  or  Framboesia  tropica  ...  ...  ...  ...  ...  632 

C.  Syphilis           ...             ...  ...  ...  ...  ..'.  ...  632 

D.  Bronchial  Spirochsetosis  ...  ...  ...  ...  ...  632 

VI. — Malaria             ...            ...  ...  ...  ...  ...  ...  633 

VII.— Balantidian  Dysentery  ...  ...  ...  ...  ...  637 

A        Plathelminthes  (Flat  Worms)          ...  ...  ...  ...  ...  ...  638 

Fascioliasis               ...            ...  ...  ...  -...  ...  ...  638 

Fasciola  hepatica  ...             ...  ...  ...  ...  ...  ...  638 

Fasciolopsis  buski ...             ...  ...  ...  ...  ...  ...  638 

Paragonimiasis         ...            ...  ...  ...  ...  ...  ...  639 

Par agonimus  ringer i           ...  ...  ...  ...  ...  ...  639 

Clonorchis  sinensis                ...  ...  ...  ...  ...  ...  640 

BiLHARZiASis               ...             ...  ...  ...  ...  ...  ...  641 

Schistosoma  hamatobium     ...  ...  ...  ...  ..'.  ...  641 

Cestodes      ...            ...            ...  ...  ...  ...  ...  ...  644 

General   ...             ...             ...  ...  ...  ...  ...  ...  644 

Dibothriocephalus  latus        ...  ...  ...  ...  ...  ...  658 

Sparganum  mansoni            ...  ...  ...  ...  ...  ...  659 

Dipylidium  canimun  {Tania  cucuinerind)       ...  ...  ...  ...  659 

Hymenolepis  nana                ...  ...  ...  ...  ...  ...  661 

Tcenia  solium         ...             ...  ...  ...  ...  ...  ...  662 

T(^nia  saginata      ...             ...  ...  ...  ...  ...  ...  667 

Nematodes  ...            ...            ...  ...  ...  ...  ...  ...  674 

Strong}' loides  steiroralis       ...  ...  ...  ...  ...  ..  674 

Dracuncu/us  7nedinensis  {DiSiContiasis)  ...  ,,.  ...  675 

Filaria  bancrofli  ...              ...  ...  ...  ...  ...  ...  676 

Loa  /oa    ...             ...             ...  ...  ...  ...  ...  ...  678 

TricJniris  trichiura               ...  ...  ...  ...  ...  ...  678 

Trichinella  spiralis               ...  ...  ...  ...  ...  ...  680 

Eustrcngylus  gigas               ...              ...  ...  ...              ...  ...  681 

Ancylostoina  duodenale  (Ancylostomiasis)  ...  ...  ...  ...  682 

Ascaris  lumbricoides  (Ascariasis)  ...  ...  ...             ...  ...  687 

6?j»y«m  ew7«?V7//am  (Oxyuriasis)     ...  ...  ...             ...  ...  694 


CONTENTS  xxi 


So  PPLEMENT— <r^;//^/. 

Hirudinea  (Leeches) 
Arthropoda 

Arachnoidea 

Leptus  aiituimialis  (Grass,  Harvest,  or  Gooseberry  Mite) 
Kedani,  Akaneesch  (The  Japanese  River  or  Inundation  Disease) 
Dernianyssiis  gallince  {avitwi) 
Ixodes  reduvhis  {ricinus) 
Sarcopfes  scadi'ei  {Scabies)    ... 
Demodex  folliculorutn  ...  ... 

Deniodex  follicu  lor  urn  canis... 
Insecta 

Pediculus  capitis  ( Head  Louse)  ...  ...  ... 

Pedictilus  vestivienii  {Q\o\}cie%  l^QMiSe) 

Phthirius  inguinalis  {Pediculus pubis)  (Crab  Louse)'      ... 

Cimex  {Acanthia)  lectularia  {Cimex  leciularius)  (BedlBug) 

Pulex  irritatis  {)^\xvc\2Xi  Y\&a) 

Derniatophilus  {Sarcopsylla)  penetrans  (Sand  Flea) 

Myiasis  ... 

Myiasis  externa     ... 

Gastricolous  Oestridce  (Creeping  Disease) 


APPENDIX  ON  PROTOZOOLOGY  ... 

L— Notes  on  Recent  Researches 

Differences  between  Entamoeba  histolytica  and  E.  coli 

Phagedaenic  Amoebae    ... 

Endamceba gingivalis   ... 

Entama'ba  kartulisi 

Cra/^2«  and  Craigiasis  .. . 

Human  Trichomoniasis 

Chiloniasiix  {Tetratnitus)  mesnili 

Giardia  {Lamblia)  intestinalis    ... 

Cercomonas  hotninis 

Transmissive  Phase  of  Trypanosomes  in  Vertebrates 

Trypanosoma  lewisi 

Blepharoplastless  Trypanosomes 

The   Experimental    Introduction    of    certain   Insect   Flagellates 
Vertebrates,  and  its  bearing  on  the  Evolution  of  Leishmaniasis 

The  Tr?insmh?,\ox\  oi  Spirockata  dtittoni  ... 

Spirochata  bronchialis   .. 

The  Spirochtetes  of  the  Human  Mouth     ... 

Coccidia  in  Cattle 

The  Haemosporidia 

The  Leucocytozoa  of  Birds 
II. — Formula  of  some  Culture  Media  ... 

Culture  Media  for  growing  Amoebae 

Culture  Media  for  the  growth  of  Protozoa  parasitic  in  the  Blood 
III. — Brief  Notes  on  General  Protozoological  Technique 

Fresh  Material 

Stained  Material 

Fixatives 

Stains 


699 
702 
702 
702 
703 

704 
704 
708 
709 
709 
709 
710 
711 
7>3 
714 
714 
715 
715 
729 


733 

733 

•  • 

733 

.. 

733 

.. 

734 

734 

734 

735 

736 

736 

737 

377 

737 

various 

737 

739 

739 

740 

741 

742 

742 

742 

742 

744 

745 

745 

747 

748 

749 

XXI 1 


THE   ANIMAL   PARASITES   OF   MAN 


APPENDIX  ON  TREMATODA  AND  NEMATODA 

...     753 

Trematoda 

...     753 

Artyfechinostomum  sufrartyfex  ... 

...     753 

Metagoniimis  ( Yokogawd)  yokogawai 

...     753 

Opisthorchis  sp. 

...     753 

Schistosome  cercarim 

...    753 

Distomata  cercariiF 

-    753 

Group.     Ferrocercous  cercarice    ... 

•  ••    753 

Family.     Schistosoviida 

••    753 

Cercaria  bilharzia,  Leiper,  1915 

..      754 

Cercaria  dz^/iarzie//a,  heiper,  igis 

...     754 

Schistosoma  mansotii,  Sambon,  1907 

•       754 

Nematoda 

••■     754 

Ancylostomiasis 

...     754 

Ground  Itch  ... 

••     754 

Ascaris  lumbricoides     ... 

...     754 

Filariasis 

■••     755 

Onchocerca  volvtilus 

755 

Sirongyloides  stercoralis 

•••     755 

BIBLIOGRAPHY      ...            ....            ..              

...     756 

INDEX         

..     836 

LIST   OF   ILLUSTRATIONS  XXllI 


LIST    OF    ILLUSTRATIONS. 

PACK 

Fig.       I  Amcebacoli.    (After  Loesch)  ...             ...             ...             ...             ...             ...  29 

2  Encysted  intestinal  amoebae.     (After  Grassi)        ...             ...             ...             ...  31 

3  Eniama'ba  coli,\\^e-cyc\Q.     (After  Castellani  and  Chalmers)            ...             ...  32 

4  Enta7?ia'ba  CO li)  so-c&Wtdi  2m\.og2iTc\y.     (From  Minchin)      ,..             ...             ...  34 

5  Entama'ba  histolytica  {tetrageita  ioxm).     (After  Hartmann)               ...              ...  35 

6  Entamceba  histolytica,  ingestion  of  red  blood  corpuscles.     (After  Hartmann)  35 

7  Entamoeba  histolytica,  section  through  infected  intestinal  ulcer.     (After  Harris)  36 

8  Entamceba  histolytica  {tetragena),  trophozoite  and  nuclei.    (After  Hartmann)  38 

9  Entamceba  histolytica  [tetragena),  cysts.     (After  Hartmann)             ...             ...  39- 

10  Entamceba  buccalis.     (After  Leyden  and  Lowenthal)         ...             ...             ...  43 

11  Entamceba  kartuli si.     (After  Kartulis)  ...             ...             ...             ...             ...  44 

12  Amoeba  miurai.     (After  Ijima)               ...              ...              ...              ...              ...  46 

13  Chlamydophrys  enchelys.     (After  Cienkowski)     ...             ...             ...             ...  48 

14  Chlamydophrys  enchelys,  Qncy?,\.Qdi.     (After  Cienkowski)  ...             ...             ...  49 

15  Leydenia gemmipara,  Schaudinn            ...              ...              ...              ...              ...  50 

16  Trichomonas  vaginalis.     (Ater  Kunstler)             ...             ...             ...             ...  53 

17  Trichomonas  intestinalis.     (After  Grassi)             ...             ...             ..               ...  54 

18  Trichomonas  intestinalis.     (Original,  Fantham)                 ...             ...             ...  55 

19  Lamblia  intestinalis.     (After  Wenyon,  from  Minchin)      ...             ...             ...  58 

20  Lajjiblia  intestinalis.     (After  Grassi  and  Schewiakoff)      ...              ...             ...  59- 

21  Cercomonas  hominis.     (After  Davaine)                 ...             ...             ...             ...  61 

22  Cercoinonas  hominis,  ixoxu  zxi  ^c!t{vc\ozo(:.CK!i%  cy%\..     (After  Lambl)    ...             ...  61 

23  Monas pyophila.     (After  Grimm)           ...             ...             ...             ...             ...  62 

24  Prowazekia  urinaria.     (After  Sinton)                   ...              ...              .,.              ...  64 

25  Prowazekia  urinaria,  excystation.     (After  Sinton)  -          ...              ...              ...  65 

26  Trypanosotna  brucei  vci  d\\\%\on.     (After  Laveran  and  Mesnil)         ...              ...  70 

27  Trypanosoma  lewisi,  rosettes.     (After  Laveran  and  Mesnil)             ...             ...  71 

28  Trypanosoma gambiense.      (After  Dutton)             ...              ...              ...              ...  73 

29  Trypanosoma ga7nbiense,d&\Q\o^mtr\i\nvQx[ehi2^.e\\ost.    (Original,  Fantham)  73. 

30  Trypanosoma  gavibiense,^Qv^\o^vc\Qx\X.\x\  Glossina  palpalis.    (After  Robertson)  75 

31  Trypanosoma  rhodesiense.     (After  Stephens  and  Fantham)                ...              ...  77 

32  Chart  showing  daily  counts  of  number  of  Trypanosomes  per  cubic  millimetre 

of  peripheral  blood  from  a  case  of  Rhodesian  sleeping  sickness.      (After 

Ross  and  Thomson)              ...             ...             ...             ...             ...             ...  79 

33  Trypanosofna    crtizi,     schizogony.     (After     Chagas,     from     Castellani     and 

Chalmers)                ...             ...             ...             ...             ,..             ...             ...  84 

34  Trypanosoma  cruzi  in  muscle.     (After  Vianna,  from  Castellani  and  Chalmers)  85 

35  Trypanosoma    cruzi,   development    in    Triatoma   megista.      (After   Chagas, 

from  Castellani  and  Chalmers)            ...             ...              ...             ...             ..  86 

36  Trypanosoma  cruzi,   forms   found   in  salivary  glands    of    Triatoma.      (After 

Chagas,  from  Castellani  and  Chalmers)              ...              ...              ...              ...  87 

37  Trypanosoma  lewisi,  from  rat's  blood.     (After  Minchin)    ...              ...              ...  89 

38  Trypanosoma  lewisi,  {xovci  %'(.ovc\2i.c\v  oix2X-'dit2,.     (After  Minchin)      ...              ...  91 

39  Trypanosoma  lewisi,  ixom.XQ.c\.\xv(i  oi  xdX-'^Q^.     (After  Minchin)        ...              ...  92 

40  Trypaiwsoma  brucei.     (After  Laveran  and  Mesnil)             ...              ...              ...  94 

^     41  Trypanosoma  evansi.     (Original,  Fantham)         ...             ...             ...             ...  9^ 


xxiv  THE  ANIMAL  PARASITES  OF  MAN 

PAGE 

Fig.     42     Trypanosoma  eqttinum.     (After  Laveran  and  Mesnil)        ...             ...             ...  96 

43  Trypanosoma  equiperdum  {(^\\g\v\-d\,Yzx\\\\9.vc{)    ...              ...              ...              ...  97 

44  Trypanosoma  theileri.     (After  Laveran  and  Mesnil)           ...              ...              ...  98 

45  Trypanosoma  vivax.     (Original,  Fanlham)           ...              ...              ...              ...  lOO 

46  Trypanosoma  congoknse.     (Original,  Fanthain)  ...              ...              ...              ...  lOO 

47  Trypanosoma  uniforme.     (Original,  Fantham)    ...              ...              ...              ...  lOO 

48  Trypanosoma  rotatoriiitn.     (After  Laveran  and  Mesnil)     ...              ...              ...  lOl 

49  Herpetomonas,  Crithidia,  Trypanosoma.     (After  Porter)  ...              ...              ...  lo.^ 

50  Leishmania  donovani.      (After  Christophers,  Patton,  Leishman ;  from  Cas- 

tellani  and  Chalmers)           ...             ...             ...             ...             ...             ...  106 

51  Toxoplasma  gondii.     (After  Laveran  and  MaruUaz,  from  7><7/.  Z)w.  ^«//(i?/««)  1 13 

52  Toxoplasma  pyrogenes.     (After  Ca.ste\\a.ni,  from  Tr op.  Dis.  Bulletin)             ...  1 13 

53  Spirochcetabalbianii.     (After  Fantham  and  Porter)            ...              ...              ...  1 14 

54  Spirochceta  duttoni.     (After  Fantham)  ...              ...              ...              ...              ...  1 17 

55  SpirochcEta  duttoni  and  its  coccoid  bodies  in  the  tick.     (After  Fantham)        ...  118 

56  Treponema  pallidum.     (After  Bell,  from  Ca«;tellani  and  Chalmers)...             ..  124 

57  Treponema  pallidum,  2i\iip2iX2L\.\x%  iox  c\x\t\w2ii\on  o{.     (After  Noguchi)              ...  125 

58  Treponema pertenue.     (After  Castellani  and  Chalmers)      ...             ...             ...  127 

59  Monocystis  agilis.     (After  Stein)            ...             ...             ...             ...             ...  130 

60  Cr^^arma /tfw^a,  stages  of  growth  of  trophozoite                ...             ...             ...  130 

61  Xyphorhynchus Jirvms.     (After  Leger)                 ...              ...              ...              ...  13 1 

62  Gregarina  munieri.     (After  Schewiakoff)             ...              ...              ...              ...  131 

63  Monocystis  agilis,  si^oxQ?,.     (After  Biitschli)         ...             ...             ...             ...  132 

64  Gregarines,  conjugation  and  spore  formation.     (After  Calkins  and  Siedlecki, 

modified)                ...             ...             ...             ...             ...             ...             ...  133 

65  Stylorhynchus  oblongatus.  cy?,X.  2LX\di  gz^meies.     (After  Leger)             ...             ...  133 

66  Gregarines,  various  spores.     (After  Leger)           ...              ...              ...              ...  134 

67  Eimeria  [Coccidium)  schuhergi,  life-cycle  diagram  of.     (After  Schaudinn)    ...  139 

68  Eimeria  avium  in  gut  epithelium  of  grouse  chick.     (After  Fantham)              ..  143 

69  ^zw^^/d!  az^/«w,  life-cycle,  diagram  of.     (After  Fantham)                ...              ...  144 

70  iS'zw^r/dt  j/?<?a'(«  in  section  of  rabbit's  intestine       ...              ...              ...              ...  145 

71  ^m^rza  j/?W^,  oocysts  from  rabbit's  liver.     (After  Leuckart)         ...              ...  146 

72  Eimeria  stieda,  ST^oxts,.     (After  Balbiani)              ...              ...              ...              ...  146 

73  Eimeria  stied(P,  %c\\\zogor\y.     (After  R.  Pfeififer)                  ...              ...              ...  146 

74  ^?w«r?'rt;  5/?V</(?,  section  through  infected  nodule  of  liver   ...              ...              ...  147 

75  Isospora  bigemina.     (After  Sti'es)           ...              ...              ...              ...              ..  150 

76  Hcemoproteus  {Halteridium)  columbce,  life-cycle.     (After   Aragao,  from  Cas- 

tellani and  Chalmers)           ...             ...             ...             ...             ...             ...  152 

77  Leucocytozo'dn  lovali.     (After  Fantham)                  ...              ...              ...              ...  153 

78  Hsemogregarines  from  lizards.      (After  Fran9a)    ...              ...              ...              ...  154 

79  Leucocytogregarina  canis,  life-cycle.    (After  Chastophers,  from  Castellani  and 

Chalmers)               ...             ...             ...             ...             ...             ...  155 

80  Plasmodium  vivax,  \\fe-cyc\e.     (Afier  Schaudinn  and  Grassi)           ...              ...  160 

81  Malignant  tertian  malarial  parasite  in  intestine  of  Anopheles.     (After  Grassi)  162 

82  Ookinete   of  malignant    tertian    malaria    in    stomach    of  Atiopheles.      (After 

Grassi)     ...             ...             ...             ...             ...             ...             ...             ...  162 

83  Section  of  stomach  of  ^;/^/,^*?/^j- with  malarial  oocysts.     (After  Grassi)          ...  163 

84  Sporulation  of  malarial  parasites  in  ^//^//^^/,?j-.     (After  Grassi)        ...              ...  163 

85  Tertian  malarial  parasite  in  human  red  blood  corpuscles.      (After  Mannaberg)  165 

86  Quartan  malarial  parasite  in  human  red  corpuscles.     (After  Manson)               ...  166 

87  Malignant  malarial  parasite  in  human  red  corpuscles.     (After  Manson)          ...  168 

88  Malarial  crescents.     (After  Mannaberg)                ...              ...              ...              ...  168 

Section  through  tubule  of  salivary  gland  of  Anopheles  infected  with  malarial 

sporozoites.     (After  Grassi)                  ...              ...              ...              ...              ...  169 

■90     Nuttallia  equi,    life-cycle   in    red   blood    corpuscles.       (After    Nuttall    and 

Strickland)             ...             ...             ...             ...             ...             ...             ...  173 


LIST  OF  ILLUSTRATIONS 


XXV 


PAGE 


Fig. 


91 

92 
93 
94 
95 
96 

97 
98 

99 

ICO 

101 

102 
103 
104 

105 
106 
107 
108 
109 
1 10 
III 
112 

1^3 
114 

"S 
116 
117 
118 
119 
120 
121 
122 
123 
124 

125 
126 
127 
128 
129 
130 

131 
132 

133 
134 
13s 
136 
137 

138 
139 
140 
141 
142 

143 
144 


Babesia  {Piroplasnia)  canis^  life-cycle  in  blood  of  dog.     (After  Nuttall  and 

Graham-Smith) 
Theileria  parva.     (After  Nuttall  and  Fantham) 

Myxosporidian  spores  and  infected  gill  of  fish.     (After  J.  Mliller)  ... 
Myxobolus  jnulleri,  spore.     (After  Biitschli) 
MyxoboluSy  %chtvc\2i  oi  s\>oxQ.     (After  Doflein)     ... 
Chloromyxum  leydigi.     (After  Thelohan) 

Myxobolus  ffeifferi,  spore  formation.     (After  Keysselitz,  from  Minchin) 
Nosema  apis.     (After  Fantham  and  Porter) 
Nosema  bombycis  from  silkworm.     (After  Balbiani) 
Nosema  bombycis ^  spores.     (After  Thelohan) 
Hexactinomyxon  psam/noryctis,  spore.     (After  Stole) 

•  Sarcocystis  miescheriana  in  muscle  of  pig.     (After  Kiihn)  "... 

Sarcocystis  miescheriana,  mature  trophozoite 

Sarcocystis  tenella  in  section,  as  seen  in  oesophagus  of  sheep 

Sarcocystis  teneUa^yowxi^  Uo'^YiOzoxit..     (After  Bertram)  ... 

Sarcocystis  miescheriana,  end  portion  of  trophozoite.     (After  Bertram) 

Sarcocystis  blanchardi  from  ox.     (From  Wasielewski,  after  van  Eecke) 

Sarcocystis  tenella.     (After  Laveran  and  Mesnil) 

Haplosporidium  heterocirri.     (After  CauUery  and  Mesnil) 

Haplosporidian  spores.     (After  Caullery  and  Mesnil) 

Rhinosporidium  kinealyi,  portion  of  ripe  cyst.     (After  Minchin  and  Fantham) 

Balantidium  coli.     (After  Leuckart)     ... 

Balantidium  coli,  free  and  encysted.     (After  Casagrandi  and  Barbagallo 

Balantidium  minutum.     (After  Schaudinn) 

Nyctotherus faha.     (After  Schaudinn)  ... 

Nyctotherus  giganteus.     (After  Krause) 

Nyctotherus  africantcs.     (After  Castellani) 

Trachoma  bodies  in  conjunctival  cells.     (Original,  Fantham) 

Half  of  a  transverse  section  through  Fasciola  hepatica,  L. 

Harmostomtcm  leptostomum,  Olss. 

Median  section  through  the  anterior  part  of  Fasciola  hepatica 

Polystomum  integerrimum.     (After  Zeller) 

Allocreadiur/i  isoporum,'Loosa.     (After  Looss)  ... 

Terminal  flame  cell  of  the  excretory  system.     (Stephens).,. 

Diagram  of  female  genitalia.     (Stephens) 

Diagram  of  male  and  part  of  female  genitalia.     (Stephens) 

Osnm  o{  Fasciola  hepatica,  1^. 

Mira.cid\\im  of  Fasciola  he/>atica.     (After  Leuckart) 

A  group  of  cercariae  of  Echinostoma  sp.  ...      > 

Development  of  Fasciola  hepatica,  L.     (After  Leuckart)... 

Young  redia  oi  Fasciola  hepatica.     (From  Leuckart) 

Older  redia.  of  jDistoma  echinatum 

Cercana.  of  Fasciola  hepatica.     (After  Leuckart) 

Encysted  cercaria  of  Fasciola  hepatica.     (After  Leuckart) 

Watsonius  watsoni.     (After  Shipley)    ... 

Watsonius  watsoni :  ventral  projection  composed  from  a  series  of 

sections.     (After  Stiles  and  Goldberger) 
Gastrodiscus  hominis.     (After  Leuckart) 
Fasciola  hepatica,  L. 

Fasciola  hepatica,  showing  the  gut  and  its  branches 
Fasciola  hepatica,  \,.     (After  Glaus)     ... 
Fasciola  hepatica  :  egg  from  liver  of  sheep.     (After  Thomas) 
Limnceus  truncatulus,M\\\\.     (From  Leuckart)... 
Yowng  Fasciola  hepatica.     (From  Leuckart) 


XXvi  THE  ANIMAL  PARASITES  OF  MAN 

PAGE 

Fig,    145  Fasciola gigantica.     (After  Looss)         ...             ...             •••  •••  •••  243 

146  Fasciolopsis  buski,  \.2iXi\i.     (After  Odhner)          ...             ...  •••  •••  245 

147  Fasciolopsis  raihouisi,  VoU.     (After  Claus)          ...              ...  •••  •••  246 

148  Fasciolopsis fiillebo7-ni.     (After  Fulleborn)          ...             ...  •••  ...  248 

149  Paragonimus  ringeri,  Cohh.     (After  Katsurada)                ...  ...  .••  250 

150  Paragonimus  ringeri,  Cohh.     (After  Kubo)        ...              ...  ...  ••■  250 

150A     Paragonimus  westermanii,  Ktxh.     (After  Leuckart)       ...  ...  ...  25a 

15 '  "Eggoi  Paragonimus  ringeri,  Cohh.     (After  Katsurada)  ...  ...  ...  251 

^52  "Egg  o{  Opisthoj'c his felineus    ...              ...              ...              .••  •••  •••  253 

153  Opisthorchis felineus.     (After  Stiles  and  Hassall)               ...'  ...  ...  253 

154  Opisthorchis pseudofelineus.     (After  Stiles)          ...             ...  .-•  ...  254 

155  Parapisthorchis  caninus.     (After  Stephens)         ...             ...  •••  ...  256 

156  Amphiinerus  nove}'ca,Bxz.Mn.     (After  McConnell)             ...  ...  ..  257 

157  Metorchis  conjunctus.     (After  Cobbold)                ...             ...  ■••  .••  258 

158  Clonorc his  sinensis.     (After  Looss)       ...             ...             ...  .••  ...  259 

1$}  OwSioi  Clonorchis  sinensis.     (After  Looss)          ...             ...  ...  ...  259 

160  CJonorchis  endemicus.     (After  Looss)  ...              ...              ...  .  .  .••  260 

161  Clonorchis  endemicus :  tggs.     (After  Looss)        ...             ...  ...  ...  260 

162  Metorchis  truncalus ...             ...             ...             ...             •.■  ••.  •••  262 

163  Heterophyes  heterophyes.     (After  Looss)               ...             ...  ...  ...  263 

164  MetagonijHUs yokogawai.     (After  Leiper)            ...             ...  ...  ...  264 

165  Dicrocceliu7n  dendriticum         ...              ...              ...              .••  ...  ...  265 

166  Eggs  of  Dicroccelium  dendriticum         ...             ...             ...  ...  ...  266 

167  M\xsLC\d\2i.o{ Dicroccelium  dendriticum.     (After  Leuckart)  ...  ...  266 

168  Echinostoma  ilocanum.     (After  Brumpt)               ...              ...  ...  ...  268 

169  Echinostoma  ilocanum.     (After  Leiper)               ...             ...  ...  ...  268 

170  Echinostoma  malayanumyL.Q\^ex.     (After  Leiper)             ...  ...  ...  269 

171  Schistosoma  hismatobium.     (After  Looss)              ...              ...  ...  ...  270 

172  Transverse  section  through   a   pair  of  Schistosoma  hcematobiutn  in  copula. 

(After  Leuckart)    ...             ...             ...             ...             ..•  ...  ...  271 

173  Anterior  end  of  the  male  Schistosoma  hcematobiuvt.     (After  Looss)  ...  271 

174  Schistosoma  hcematobium.     (After  Leuckart)        ...             ...  ...  ...  276 

175  Schistosoma  hcematobiu7Jt,  owxvci  oL     (After  Looss)             ...  ...  ...  277 

176  Schistosoma Japonicum.     (After  Katsurada)         ...              ...  ...  ...  278 

177  Schistosojna  japonicum.     (After  Katsurada)         ...             ...  ...  ...  279 

178  Schistosojna  japonicum.     (After  Looss)                 ...             ...  ...  ...  279 

\%o\  Schistosoma  japonicum  ixoxciAog.     (After  Katsurada)         ...  ...  ...  280 

181  J 

182  Schistosoma  japonicum.     (After  Catto)                  ...             ...  ...  ...  281 

183  Schistosoma  japonicum.     (After  Katsurada)         ...             ...  ...  ...  282 

184  Schematic  representation  of  a  small  part  of  a  transverse  section  of  Ligula  sp. 

(After  Blochmann)                ...             ...             ...             ...  ...  ...  287 

185  Half  of  a  transverse  section  through  a  proglottis  of  Tcenia  crassicollis  ...  288 

186  Dipylidium  (anintim.     (After  Benham)               ...              ..  ...  ...  289 

187  Longitudinal  section  of  the  head  and  neck  of  Tcenia  crassicollis  ...  ...  290 

188  Tcenia  ccenurus.     (After  Niemisec)        ...              ...              ...  ...  ...  291 

189  Yo\xx\g  Acanthobothrium  coronatnm.     (After  Pintner)        ...  ...  ...  292 

190  Scolex  of  a  cysticercoid  from  Arion  sp.     (After  Pintner)  ...  ...  ...  292 

191  Proglottis  of  7>«z'a  .y<i^/«fl!/a,  Goeze,  showing  genitalia    ...  ...  ...  293 

192  Dibothriocephalus  lattis.     (After  Benham  and  Sommer  and  Landois)  ...  294 

193  Diagram  of  genitalia  of  a  Cestode.     (Stephens)                   ...  ...  ...  295 

194  Part  of  a  transverse  section  through  a  proglottis  of  Dibothriocephalus  latus  ...  296 

195  Y^ggoi  Diplog07ioporus grandis.     (After  Kurimoto)             ...  ...  ...  298 

196  \}\.tx\vi^  &gg  oi  Tania  saginata.     (After  Leuckart)             ...  ...  ...  298 

197  Oncosphere  of  Tcenia  a/ricana  (after  v.  Linstow)  and  oncosphere  of  Dipy- 

lidium caninum.     (After  Grassi  and  Rovelli)  ...              ...  ...  ...  299 


LIST  OF  ILLUSTRATIONS  XXVll 

PAGE 

Fig.   198  Diagram  of  a  cysticercoid.     (Stephens)  ...  ...  ...  ...     301 

199  Diagram  of  a  cysticercus.     (Stephens)  ...  ...  ...  ...     301 

200  Diagram  of  development  of  a  cysticercus.     (Stephens)       ...  ...  ...     303 

201  Section  through  a.  piece  of  a.  Ccenuf'Ms  ceredrah's     ...  ...  ...  ...     304 

202  Median  section  through  a  cysticercus.     (After  Leuckart)  ...  ...  ...     304 

203  Qys/t'cercus pisi/ormis  in  a.T\  evaginated  condition  ...  ...  ...     304 

204  Various  chains  of  segments  of  Z)/<^<7/'^r/<?<r<?//^a/«5- /a/MJ-      ...  ...  ...     311 

205  Transverse  section  of  the  head  of  £)ii>o^/irzocepka/us /attts  ...  ...     311 

206  Fairly  mature  proglottis  of  Z)z<^c//zrz£>^^//^a/?^j /a/M^  ...  ...  ...     311 

207  Dibothriocephahis  lahis,     (After  Benham  and  Schauinsland)  ...  ...     312 

208  VXerocercoid  of  Dibothriocephalus  latus  ...  ...  ...  ...     313 

209  A  piece  of  the  body  wall  of  the  Burbot,  Z^/d!  zv//^aw      ...  ...  ...     313 

210  CephaWc  end  of  Dibothriocephalus  cordatHs.     (After  Leuckart)        ...  ...     315 

211  Diplogonoporus grandisy\X\^ey  1899.     (After  Ijiraa  and  Kurimoto)  ...     317 

212  Diplogonoporus grandis.     (After  Ijima  and  Kurimoto)      ...  ...  ...     317 

213  Cephalic  end  of  ^^^fl^-^awww  ;//a«j-^«/,  Cobb.     (After  Leuckart)      ...  ...     318 

214  Sparganuin  mansoni .     (After  Ijima  and  Murata)  ...  ...  ...     318 

215  Sparganum  pro  lifer.     (After  Ijima)      ...  ...  ...  ...  •••319 

216  Spai'ganum prolifefitm.     (After  Stiles)  ...  ...  ...  ...     319 

217  Dipyliditim  caninuiu.     (After  Diamarr)  ...  ...  ...  ...     320 

218  Dipylidium  caninum.     (After  Benham  and  Moniez)         ...  ...  ...     320 

219  Dipylidium  caninum  :  central  portion  of  a  proglottis.     (After  Neumann  and 

Railliet)  ... 

220  Dipylidium  caninum  :  development  of  embryo.     (After  Benham,  Grassi,  and 

Rovelli)  ... 

221  Larva  (cysticercoid)  of  Dipylidium  caninum.     (After  Grassi  and  Rovelli) 

222  Hymenolepis  nana,  v.  Sieb.     (After  Leuckart)    ... 

223  Hymenolepis  nana:  head.     (After  Mertens) 

224  Hymenolepis  nana  :  an  egg.     (After  Grassi) 

225  Longitudinal  section  through  the  intestinal  villus  of  a  rat.     (After  Grassi  and 

Rovelli)  ... 

226  Hymenolepis  nana  {murina)  :    cross-section  of  proglottis  from  a  rat.     (After 

v.  Linstow) 

227  Hymenolepis  nana :   longitudinal  section  of  an  embryo.     (After  Grassi  and 

Rovelli)  ... 

228  Hytnenolepis  di^ninuta.     (After  Zschokke) 

229  Hymenolepis  diminuta.     (After  Grassi) 

230  Hymenolepis  diminuta.     (After  Bizzozero) 

231  Hymenolepis  diminuta.     (Stephens,  after  NicoU  and  Minchin) 

232  Hymenolepis  lanceolata.     (After  Krabbe) 

233  Hymenolepis  lanceolata.     (After  Wolffhiigel) 

234  ScoXeyi  of  Davainea  madagascariensis.     (After  Blanchard) 

235  Two  fairly  mature  proglottids  of  7<2;/za  WzV/w  ... 

236  WeaA  of  Tcenia  solium 

237  Large  and  small  hooks  of  Tcenia  solium.     (After  Leuckart) 

238  TcEtiia  soliiwi.     (After  Leuckart) 

239  Two  mature  proglottids  of  7}2«/a  Wzz/w 

240  Large  and  small  booklets  of  Tctnia  marginaia.     (After  Leuckart)  ... 

241  yfa.twxe  segment  of  TiEnia  saginata 

242  Cephalic  end  of  Tcenia  saginata 

243  Tania  saginata.     (After  Leuckart) 

244  A  piece  of  the  muscle  of  the  ox,  with  three  specimens  of  Cysticercus  bovis 

(After  Ostertag)  ... 

245  "^.a-twresegrnent  of  Tcenia  africana.     (After  v.  Linstow)  ... 

246  Proglottis  of  Tenia  africana.     (After  v.  Linstow) 

247  Head  of  Tcenia  africana.     (After  v.  Linstow)     ... 


XXVlll  THE  ANIMAL  PARASITES  OF  MAN 

PAGE 

Fig.  248     Tania  confiisa.     (After  Guyer)  ...  ...  ...  ...  ...  344 

249  Tania  confusa.     (After  Ward)  ...  ...  ...  ...  ...  344 

250  Tcenia  echinococcus    ...  ...  ...  ...  ...  ...  ...  345 

251  Echinococcus  veterinorw)i.     (After  Leuckart)      ...  ...  ...  ...  347 

^      r  Diagrams  of  mode  of  formation  of  brood  capsule  and  scolices  (Stephens)     ...  348 

253  Section  through  an  invaginated  echinococcus  scolex.     (After  Deve)  ...  350 

254  A  piece  of  the  wall  of  an  Echinococcus  veterinoriivi  stretched  out  and  seen  from 

the  internal  surface  ...  ...  ...  ...  ...  •••  35° 

255  Echinococcus  hominis  in  the  liver.     (After  Ostertag,  from  Thomas)  ,..  351 

256  Section  through  an  echinococcus  scolex  in  process  of  vesicular  metamorphosis. 

(After  Deve)         ...  ...  ...  ...  ...  ...  ...  35^ 

^^'     \  Diagram  of  transformation  of  a  scolex  into  a  daughter  cyst.     (Stephens)      ...  352 
257A ) 

258  Hooklets  of  echinococcus.     (After  Leuckart)      ...  ..  ...  ...  355 

259  Echinococcus  multilocularis  \n\\\^\w^x  o{\\i^  o\.     (After  Ostertag)  ...  357 

260  Diagram  of  a  transverse  section  of  .(^j^flr^V /MW(5rzV^?V/(fj.     (After  Brandes)    ...  362 

261  Anlerior  end  of  an  Ascaris  megahce/>ha/a.     (After  Nassonow)         ...  ...  362 

262  Transverse  section  through  Ascaris  lumbricoides  at  the  level  of  the  oesophagus 

behind  the  nerve  ring.     (After  Goldschmidt)  ...  ...  ...  .■•  3^4 

263  Schematic  representation  of  the  nervous  system  of  a  male  Ascaris  megalo- 

cephala.     (After  Brandes)    ...  ...  ...  ...  ...  ...  3^5 

264  Diagram  of  female  genitalia    ...  ..  ...  ...  ...  ...  3^8 

264A  Diagram  of  male  genitalia  of  a  strongylid  ...  ...  ...  ...  368 

265  Transverse  section  through  the  ovarian  tube  of  Belascaris  cati  of  the  cat         ...  369 

266  lA'sXo.  Q{\!nex\\2}o^\\\z  ioxvn  oi  Angiostonmin  nigrovenosuiii  ...  ...  370 

267  Transverse  section  through  the  posterior  extremity  of  the  body  of  Ascaris 

lumbricoides  {maXe)  ...  ...  ...  ...  ...  •••  37^ 

268  Hind  end  of  a  male  Ascaris  lumbricoides  cut  across  at  the  level  of  the  dilator 

cells  of  the  gut.     (After  Goldschmidt)  ...  ...  ...  ...  371 

269  A  piece  of  the  trunk  muscle  of  the  pig  with  encapsuled  embryonic  Trichinge...  373 

270  Strongyloides  siercoralis,  itmSiU.     (After  Looss)  ...  ...  ...  380 

271  Strongyloides  siercoralis,  mzXe.     (After  Looss)    ...  ...  ...  ...  380 

272  Strongyloides  stercoralis,iexfX2\e.     (After  Loo.ss)...  ...  ...  ...  382 

273  Strongyloides  stercoralis.     (After  Looss)  ...  ...  ...  ...  3^2 

274  Strongyloides  stercoralis.     (After  Looss)  ...  ...  ...  ...  3^3 

275  Gnathostoma  siamense.     (After  Levinsen)  ...  ...  ...  ...  385 

276  Gmnedi  worm  {Dracuncul us  medinensis).     (After  Leuckart)  ...  ...  387 

277  Anterior  extremity  of  Guinea  worm.     (After  Leuckart)     ...  ...  ...  387 

278  Dracunculus  medinensis.     (After  Claus)  ...  ...  ...  ..'.  3^7 

279  Transverse  section  of  female  Guinea  worm.     (After  Leuckart)         ...  ...  388 

280  Cyclo/>s  virescens,  (emsde  ...  ...  ...  ...  ...  ...  3^9 

281  Filaria  bancrofti.     (After  Leiper)  ...  ...  ...  ...  ...  39^^ 

282  Mf.  bancrofti  in    thick  film,    dried    and    stained    with  hsematoxylin.     (After 

Fiilleborn)  ...  ...  ...  ...  ...  ...  ...  397 

283  Schematic  drawings  of  the  anatomy  of  Ml.   loa  and  Mf.   bancrofti.     (After 

Fiilleborn)  ...  ...  ...  ...  ...  ...  ...  399 

284  F.  demarquayi.     (After  Leiper)  ...  ...  ...  ...  ...  4^3 

285  Mf.  demarquayi  \n  thick  film,  dried  and  stained  with  haematoxylin.     (After 

Fiilleborn)  ...  ...  ...  ...  ...  ...  ...  4^4 

286  Filaria  {Tj  conjunctivie.     (After  Addario)  ...  ...  ..  ...  405 

287  Filaria  (?)  conJunclivcE.     (After  Grassi)  ...  ...  ...  ...  405 

288  Setaria  equina.     (After  Railliet)  ...  ...  ...  ...  ...  408 

289  Setaria  equina  :  anterior  end.     (After  Railliet)  ...  ...  ...  ...  408 

290  Loa  loa  :  the  anterior  end  of  the  male.     (After  R.  Blanchard)         ...  ...  410 

291  Zca /^a  ;  anterior  portion  of  the  female.     (After  Looss)    ...  ...  ...  410 


LIST  OF  ILLUSTRATIONS  XXIX 

PAGE 

Fig.  292     Loa  loa  in  situ.     (After  Flilleborn  and  Rodenwaldt)           ...             ...  ...  410 

293  Loa  loa :  male  and  female.     (After  Looss)           ...             ...             ...  ...  410 

294  Loa  loa:  the  hind  end  of  a  male  and  of  a  female.     (After  Looss)  ...  ...  411 

295  Loa  loa  :  lateral  view  of  tail  of  male  showing  papillae.    (After  Lane  and  Leiper)  411 

296  Loa  loa.     (After  Leiper)          ...             ...             ...             ...             ...  ...  411 

297  i^. /m  :  in  thick  film,  dried  and  stained  with  hsematoxylin.    (After  Fiilleborn)  413 

298  Acanthocheilonema Persians.     (After  Leiper)       ...             ...             ...  ...  414 

299  Mf.  ferstans.     (After  Flilleborn)           ...             ...             ...             ...  ...  415 

300  Dirofilaria  magalhdesi.     (After  v.  Linstow)        ...             ...             ...  ...  417 

301  Trichuris  trichiura  ...              ...              ...              ...              ...              ...  ...  420 

302  Trichinella  spiralis.     (After  Claus)      ...             ...             ...             ...  ...  422 

303  Isolated  muscular  fibre  of  a  rat,  invaded  by  Trichinella.     (After  Hertwig- 

Graham)...              ...              ...              ...              ...              ...              ...  ...  425 

304  Calcified  Trichinella  in  the  muscular  system  of  a  pig.     (After  Ostertag)        ...  426 

305  Various  phases  of  the  calcification  of  Trichinella  of  the  muscles      ...  ...  426 

306  Diociophyme gigas.     (After  Railliet)     ...             ...             ...             ...  ...  432 

307  'E'ggs  oi  Dioctophy77te  gigas.     (After  Railliet)      ...             ...             ...  ...  432 

308  Metastrongylus  apri.     (Stephens)          ...             ...             ...             ...  ...  433 

I  Irichostrongyhis  instabilis.     (After  Looss)         ...             ...             ...  ...  434 

^       I  Trichostrongylus probolurus.     (After  Looss)        ...             ...             ...  ...  435 

3  3  \  Trichostrongylus  vitrinus.     (After  Looss)            ...             ...             ...  ...  436 

3H  i 

3^5  \  Hcemonchus  contortus.     (After  Ransom)               ...             ...             ...  ...  437 

316  [ 

317  Mecistocirrus fordi.     (After  Stephens)  ...             ...             ...             ...  ...  439 

318  Ternidens  deminuius.     (After  Railliet  and  Henry)             ...             ...  ..  440 

\CEsophagostoviumstephanostomiim\2Ji.thomasi.     (After  Thomas)  ...  ...  442 

r  CEsophagostomum  stephanostomnm  var.  thomasi.     (After  Thomas)...  ...  444 

^^■^  i  Ancylosloma  duode7iale,  male  a.nd  (emsile.     (After  Looss)                  ...  ...  446 

324  ) 

325  Ancylostoma  duodenale,  showing  ventral  teeth.     (After  Looss)         ...  ...  447 

326  Ancylostoma  duodenale  :    diagrammatic  representation  of  excretory  system. 

(After  a  drawing  by  Looss)                 ...             ...             ...             ...  ...  448 

327  Ancylostoma  duodenale.     (After  Railliet)             ...             ...             ...  ...  449 

328  Ancylostoma  duodenale :  bursa  of  male.     (After  Looss)    ...             ...  ...  450 

329  Ancylostoma   duodenale :    eggs   in   different  stages  of  development.      (After 

Looss)      ...             ...             ...             ...             ...             ...             ...  ...  451 

330  Ancylostoma  duodenale :  larva.     (After  Leichtenstern)     ...             ...  ...  452 

331  Ancylostoma  dtwdejiale.     (After  Looss)                 ...             ...             ...  ...  453 

332  Ancylostoma  ceylanicum.     (After  Looss)               ...             ...             ...  ...  456 

333  Ancylostoma  hraziliense.     (After  Gomez  de  Faria)             ...             ...  ...  456 

334  Necator  americamis.     (After  Looss)      ...             ...             ...             ...  ...  457 

335  Necator  ame7-icanus :  lateral  view.     (After  Looss)              ...             ...  ...  458 

336  Necator  americanus :  bursa  of  male.     (After  Looss)           ...             ...  ...  458 

337  Syngamus  kingi  :  anterior  end  of  male.     (After  Leiper)   ...             ...  ...  460 

338  Syngamus  kingi:  anterior  end  of  female.     (After  Leiper)               ...  ...  460 

339  'B\xx?>2i  oi  Syngamus  tj-ackealis.     (Stephens)          ...             ...             ...  ...  461 

340  Physaloptera  mordens,  luti^^eXy  igoy.     (After  Leiper)         ...             ...  ...  462 

341  Ascaris  lumbricoides.     (From  Claus)      ...              ...              ...              ...  ...  4^3 

342  0\\xxi\  o{  Ascaris  lumbricoides...              ...              ...              ...              ...  ...  4^3 

34^     Owwm  o[  Toxasca7is  limbata   ...              ...              ...              ...              ...  ...  466 

344     Transverse  section  through  the  head  pait  of  Belasca7-is  cati  from   the   cat. 

(After  Leuckart)     ...             ...             ...             ...             ...             ••■  ...  466 


Fig. 


XXX  THE  ANIMAL  PARASITES  OF  MAN 

PAGE 

^^  \  Male  a.nd  iemale  o(  Ox^uris  verf/ii'cu/aris            ...             ...  ...  ...  4^8 

347  Oxytiris  verviicularis :  egg  freshly  deposited       ...             ...  ...  •■•  4^8 

348  0:rj/wrzV  z;(?r/;^/V«/am  ;  egg  twelve  hours  after  deposition  ...  ...  ...  468 

^^^KT^ienxaXe  oi  Echinorhynchus  augustatus               ...              ...  ...  ...  47^ 

348B  Anterior  portion  of  the  female  apparatus  of  Echinorhynchus   acus.     (After 

Wagener)                ...             ...             ...             ...             ...  ...  ...  47^ 

348c  Yj^^oi  Echinorhynchus gigas.        (After  Leuckart)              ...  ...  ...  477 

348D  The  internal  organs  of  the  leech.     (After  Kennel)             ...  ...  ...  480 

348E  Hirudo  tnedicinalis,     (After  Claus)       ...              ...              ...  ...  ...  481 

349  Leptns  atitwiinaHs.     (After  Gudden)    ...             ...             ...  ...  ...  485 

350  Leptus  auhimnalis.     (After  Trouessart)               ...             ...  ...  ...  485 

351  The  kedani  mite.     (After  Tanaka)        ...             ...             ...  ...  ...  487 

352  Tetratiychus  telarius  \2i.x.  rzisseolns,\\.oc\\.     (After  Artault)  ...  ...  488 

353  Pediculoides  ventricostis.     (After  Laboulbene  and  Megnin)  ...  ...  489 

354  Nephrophages  sangutnai'his,  male,  ventral  surface.    (After  Miyake  and  Scriba)  490 

355  Nephrophages  sangtiinarius :    female,    dorsal   aspect.      (After   Miyake   and 
Scriba)    ...             ...             ...             ...             ...             ...  ...  ...  490 

356  Tydeits  inolestus.     (After  Moniez)          ...              ...              ...  ...  ...  491 

357  Dermatiyssus gallincB.     (After  Berlese)                ...             ...  ...  ...  492 

358  Dermanyssus  hirtindinis.     (After  Delafond)       ...              ...  ...  ...  492 

359  Ixodes  ricinuSy  \\\z\e.     (After  Pagenstecher)        ...             ...  ...  ...  498 

360  Yt.vi\z\eoi  Ixodes  ricinus.     (After  Pagenstecher)  ,              ...  ...  ...  498 

361  Argas  rejlextis.     (After  Pagenstecher)  ...             ...             ...  ...  ...  506 

362  Argas  per  sictis.     (After  Megnin)            ...              ...              ...  ...  ...  507 

363  Tyroglyphus  farince :  male.     (After  Berlese)       ...             ...  ...  ...  512 

364  Tyroglyphus  longior,  Gqiw.     (After  Fum.  and  Robin)       ...  ...  ...  512 

365  Rhizoglyphus  parasiticus :      male  and  female.     (After  Dalgetty)  ...  ...  514 

366  Histiogaster  [entomophagus  ?)  spermaticus.     (After  E.  Trouessart)  ...  ...  515 

367  Sarcoptes  scabiei.     (After  Fiirstenberg)...             ...             ...  ...  ...  518 

368  Sarcoptes  scabiei  :  male,  ventral  aspect.     (After  Fiirstenberg)  ...  ...  519 

369  Sarcoptes  minor  var.  cati.     (After  Raiiliet)           ...              ...  ...  ...  521 

370  Demodex folliculorum  oi  the  diOg.     (After  Megnin)            ...  ...  ...  522 

371  Lingiiatula  rhinaria :  female                ...              ...              ...  ...  ...  524 

372  \jax\z.oi  Linguatula  rhinaria  {Pentastoma  denticttlatum).  (After  Leuckart)  524 

373  linguaiula  rhinaria.     (After  M.  Koch)              ...             ...  ...  ...  525 

374  MowXh-y^zxX.^  o^  Pedictdus  vestimenti.     (After  Denny)       ...  ...  ...  533 

375  Ovum  of  the  head  louse           ...             ...             ...             ...  ...  ...  533 

376  Head  louse,  male      ...             ...             ...             ...             ...  ...  ...  533 

377  Pediculus  vestimentiy  B\xxn\.  :  adult  female         ...             ...  ...  ...  533 

378  Phthirius  inguinalis,  Leach...              ...              ...              ...  ...  ...  534 

379  Head  of  the  bed  bug  from  the  ventral  surface     ...             ...  ...  ...  535 

380  Derniatophilus penetrans  :  young  female.     (After  Moniez)  ...  ...  544 

381  Derniatophilus  penetrans :  older  female.     (After  Moniez)  ...  ...  544 

382  Pulex  irrilans          ...              ...              ...              ...              ...  ...  ...  546 

383  Larva  of  flea.     (After  Raiiliet)               ...             ...             ...  ...  ...  546 

384  Fulex  serraticeps       ...              ...              ...              ...              ...  ,,,  ...  546 

385  Head  of  a  male  and  of  a  female  Anopheles.     (After  Giles)  ...  ...  549 

386  Head  of  a  male  and  of  a  female  Culex.     (After  Giles)       ...  ...  ...  549 

387  yio^^\\i-\>2iX\%  oi  Anopheles  claviger .     (After  Grassi)            ...  ...  ...  550 

388  Anopheles  fjiaculipennis.     (Afier  Nuttall  and  Shipley)      ...  ...  ...  550 

389  Longitudinal  section  of  an   Anopheles,  showing   alimentary  canal.     (After 
Grassi)    ...             ...             ...             ...             ...             ...  ...  ...  551 

390  Anopheles  tNaculipennis,yie\gen.     (After  Grassi)               ...  ...  ,..  552 

391  'La.wa.oi  Anopheles  maculipenniSfYahx.     (After  Grassi)  ...  ...  ...  553 

392  Larva  of  Culex.     (After  Grassi)             ...             ...             ...  ...  ...  553 


LIST  OF  ILLUSTRATIONS  XXxi 

PAGE 

F^G.   393  Fupz  o(  j4nop/ieks  macu/ipenniSy  Me\g.     (Afler  Grass!)    ...             ...  ...  554 

394  Heads  of  Culex  and  Anopheles.     (After  Daniels)               ...             ...  ...  556 

395  Eggs   of  Culex,    of  Anopheles,   of  Stegomyia,    of  Taeniorhynchus,  and    of 

Psorophora             ...             ...             ...             ...             ...             ...  ...  557 

396  Diagram  showing  the  structure  of  a  typical  mosquito.     (Theobald)  ...  558 

397  Types    of    scales,    head    and    scutellar    ornamentation,    forms    of    clypeus. 

(Theobald,  etc.,  etc.)           ...              ...             ...             ...             ...  ...  559 

398  Neuration  of  wing.     Explanation  of  wing  veins  and  cells.     (Theobald)  ...  560 

399  Wmg  of  ^nophe/es  piacti/ifenmsy  Meigea            ...             ...             ...  ...  566 

4CXD  Wing  of  a  Culex        ...             ...             ...             ...             ...             ...  ...  575 

401  Wing  of  Simulium    ...             ...             ...             ...             ...             ...  ...  579 

402  Wing  of  Chironomus                ...             ...             ...             ...             ...  ...  579 

403  A  Ceratopogon,  or  midge        ...             ...             ...             ...             ...  ...  580 

404  An  owl  midge,  Phlebotonnis  sp.     (From  Giles's  "Gnats  or  Mosquitots  ")    ...  581 

405  \jax\^o{  Homaloniyia  canicular  is          ...              ...              ...              ...  ...  585 

406  'L3lx\2&  oi  Calliphora  vomitoria               ...              ...              ...              ...  ...  585 

407  Larva  of  Chrysomyia  niacellaria.     (After  Conil)                 ...              ...  ...  585 

408  The  screw-worm  fly  (Chjysomyia  macellaria)      ...              ...              ...  ...  587 

409  Ochromyia  larva  on  the  skin  of  man,  South  Africa.     (After  Blanchard)  ...  590 

410  Head  end  of  ••  larva  of  Natal."     (After  Gedoelst)              ...             ...  ...  591 

411  Lund's  larva.     (After  Gedoelst)             ...             ...              ...             ...  ...  593 

412  Dermaiobia  noxialis,  GoxxdiOi  ...              ...              ...              ...              ...  ...  597 

413  IjSiXVd^oi  Derniatobia  cyaniventris.     (After  Blanchard)     ...             ...  ...  597 

414  'L.z.xvz.of  Derviatobia  cyani7fenttis.     (After  Blanchard)     ...             ...  ...  597 

415  The  ox  gad  fly  (7a/5a«z^f  <5(7t^2««j,  Linn.)             ...             ...             ...  ...  601 

416  'Y\\t.\yi\vc\'^  {Hceinatopota  phivialis.,\Jvaxi.)            ...              ...              ...  ...  602 

417  Head  of  Glossina  longipalpis.     (Afier  Griinberg)               ...             ...  ...  604 

418  Antenna  of  Glossina  pal  iidipes,  male.     (After  Austen)      ...             ...  ..  604 

419  Glossina  palpalis  2ind  Y)\xil)yi\nxii.     (After  Brumpt)              ...              ...  ...  607 

420  The  tsetse-fly  {Glossina  morsitans^  West  wood)   ...             ...             ...  ...  608 

421  The  stinging  fly  {Stomoxys  calcitransy  Linn.)       ,..             ...             ...  ...  609 

422  Trichomonas  irom  Q.2tz\xm  2ixv^  gyxX  oi  X2X.     (Original,  Fantham)       ...  ...  735 


We  regret  to  have  taken  without  permission  from  the 
*' Transactions  of  The  Society  of  Tropical  Medicine  and 
Hygiene,"  London,  the  following  diagrams  : — 

Pages  Figures 

268        No.  169 

269        „    170 

391        „    281 

411        „    295  and  296 

414        „    298 

460        „    337  and  338 

and  tender  our  regret  to  the  Society  in  question  for  havint^ 
done  so.  ^ 


XXX  THE  ANIMAL  PARASITES  OF  MAN 

PAGE 

^  Y  Male  and  {emale  of  Oxyuri's  ver/nuularis            ...             ...  ...  ■••  4^8 

347  Oxyurts  vermicularis :  egg  freshly  deposited       ...             ...  ...  •••  468 

348  Oxyuris  vermicularis  :  egg  twelve  hours  after  deposition  ...  ...  ...  468 

T,^2>A  T'\\Qvaz\&  oi  Echinorhynchus  atigtistatus               ...              ...  ...  ...  47^ 

348B  Anterior  portion  of  the  female  apparatus  of  Echinorhynchiis   acus.     (After 

Wagener)               ...             ...             ...             ...             ...  ...  •••  47^ 

348c  'Egg  oi  Echinorhynchus gigas.        (After  Leuckart)              ...  ...  ...  477 

348D  The  internal  organs  of  the  leech.     (Afier  Kennel)             ...  ...  ...  480 

348E  Hirudo  inedicinalis.     (After  Claus)       ...              ...              ...  .  .  ...  481 

349  Leptiis  atiHiiunaHs.     (After  Gudden)    ...             ...             ...  ...  ...  485 

350  Lepttis  autuimiaJis.     (After  Trouessart)               ...             ...  ...  ...  485 

351  The  kedani  mite.     (After  Tanaka)        ...             ...             ...  ...  ...  487 

352  Tetrajiychus  telarius  va.x.  rzisseolus,\\.och.      (After  Artault)  ...  ...  488 

353  Pediadoides  ventricosiis.     (After  Laboulbene  and  Megnin)  ...  ...  489 

354  Nephrophages  sangtdnarius,  male,  ventral  surface.    (After  Miyake  and  Scriba)  490 

355  Nephrophages  sanguiriarius :    female,    dorsal    aspect.      (After    Miyake   and 

Scriba)    ...             ...             ...             ...             ...             ...  ...  ...  49° 

356  Tydeiis  molestiis.     (After  Moniez)         ...             ...             ...  ...  ...  491 

357  Dermauyssus gallincB.     (After  Berlese)                ...             ...  ...  ...  492 

358  Dermanyssus  hirundinis.     (After  Delafond)       ...              ...  ...  ...  492 

359  Ixodes  ricinus,  m2L\e.     (After  Pagenstecher)        ...             ...  ...  ...  498 

360  Femsde  oi  Ixodes  riciftus.     (After  Pagenstecher)  .              ...  ...  ...  498 

361  Argas  tejiextis.     (After  Pagenstecher)  ...             ...             ...  ...  ...  506 

362  Argas  per  sicus.     (After  Megnin)           ...             ...             ...  ...  ...  507 

363  Tyroglyphus farina :  male.     (After  Berlese)       ...             ...  ...  ...  512 

364  Tyroglyphtis  longior,  Getv.     (After  Fum.  and  Robin)       ...  ...  ...  512 

365  Rhizoglyphus  parasiticus :      male  and  female.     (After  Dalgetty)  ...  ...  514 

366  Histiogasier  {entomophagus  })  spermaticus.     (After  E.  Trouessart)  ...  ...  515 

367  Sarcoptes  scabiei.     (After  Fiirslenberg)...             ...             ...  ...  ...  518 

368  Sarcoptes  scabiei :  male,  ventral  aspect.     (After  Furstenberg)  ...  ...  519 

369  Sarcoptes  minor  sax.  cati.     (After  Raiiliet)          ...             ...  ...  ...  521 


LIST  OF  ILLUSTRATIONS  XXXI 

PAGE 

^^G-   393  Vn^pdi  oi  Anopheles  macuIipenniSf'M.ti^.     (After  Grassi)    ...             ...  ...  554 

394  Heads  of  Culex  and  Anopheles.     (After  Daniels)               ...             ...  ...  556 

395  Eggs  of  Culex,   of  Anopheles,  of  Stegomyia,    of  Taeniorhynchus,    and   of 

Psorophora             ...             ...             ...             ...             ...             ...  ...  557 

396  Diagram  showing  the  structure  of  a  typical  mosquito.     (Theobald)  ...  558 

397  Types    of    scales,    head    and    scutellar    ornamentation,    forms    of    clypeus. 

(Theobald,  etc.,  etc.)           ...             ...             ...             ...             ...  ...  559 

398  Neuration  of  wing.     Explanation  of  wing  veins  and  cells.     (Theobald)  ...  560 

399  'W'mgoi  Anopheles  niaailifemns,MQ\gen            ...             ...             ...  ...  566 

400  Wing  of  a  Culex        ...             ...             ...              ...             ...             ...  ...  575 

401  Wing  of  Simulium    ...             ...             ...              ...             ...             ...  ...  579 

402  Wing  of  Chironomus                ...             ...             ...             ...             ...  ...  579 

403  A  Ceratopogon,  or  midge        ...             ...             ...             ...             ...  ...  580 

404  An  owl  midge,  Phlebotomns  sp.     (From  Giles's  "Gnats  or  Mosquitots  ")    ...  581 

405  \j3LX\z.oi  Honialojnyia  canicularis          ...              ...              ...              ...  ...  585 

406  \jZXV2&  oi  Calliphora  vomitoria               ...              ...              ...              ...  ...  585 

407  Larva  of  Chrysomyia  viacellaria.     (After  Conil)                 ...              ...  ...  585 

408  The  screw-worm  fiy  {Chrysofnyia  macellana)      ...              ...              ...  ...  587 

409  Ochromyia  larva  on  the  skin  of  man,  South  Africa.     (After  Blanchard)  ...  590 

410  Head  end  of  **  larva  of  Natal."     (After  Gedoelst)              ...             ...  ...  591 

411  Lund's  larva.     (After  Gedoelst)             ...             ...             ...             ...  ...  593 

412  Der77iatobia  noxialis,  Goudot  ...              ...              ...              ...              ...  ...  597 

413  'L^rwz.  o{  Dermatohia  cyaniventris,     (After  Blanchard)     ...             ...  ...  597 

414  l^dirvQ.  oi  Derfnatobia  cyani7ientr is.     (After  Blanchard)     ...             ...  ...  597 

415  The  o\  g2id  ^y  {Tabamt^  bovimts^  Uinn.)              ...              ...              ...  ...  601 

416  The  hr\m^  {HiBniatopoia plnvialiSfUinn.)             ...              ...              ...  ...  602 

417  Hedid  oi  Glossina  longipalpis.     (After  Grunberg)               ...             ...  ...  604 

418  Antenna  of  Glossina pallidipes^  male.     (After  Austen)      ...             ...  ..  604 

419  Glossina  palpalis  zxv^  }^\x^xr\wvcv.     (After  Brumpt)             ...             ...  ...  607 

420  The  tsetse-fly  {Glossina  inorsitans.  West  wood)   ...             ...             ...  ...  608 

421  The  stinging  fly  (Stofnoxys  calcitransy  Linn.)       ...             ...              ...  ...  609 

422  7>/V^tfw<7«aj- from  caecum  and  gut  of  rat.     (Original,  Fantham)       ...  ...  735 

423  Chilomastix  {Tetramitus)  mesnili.     (Original,  Fantham)                  ...  ...  736 


XXXll  THE  ANIMAL  PARASITES   OF  MAN 


ERRATA. 


P.  31,  line  6  from  bottom:  dekte  "human,"  as  Leidy  really  worked  with  Endamceba 
blaticBy  parasitic  in  the  gut  of  the  cockroach.  ^ 

P.  43,  line  12  from  bottom  :  for  "John's  "  read  "Johns." 

P.  44,  line  13  from  hotiora:  for  ^"^  Amceba  buccalis,  Sternberg,"  read  ^^  Amoeba  buccalisy 
Steinberg." 

P.  46,  line  13  from  top  :  for  "  breath  "  read  **  breadth." 

P.  53,  In  footnote  *,  line  6  from  bottom  :  insert  "see"  before  Arch.  f.  Protistenk. 

P.  75  :  To  paragraph  regarding  development  of  the  parasite  in  the  fly's  salivary  glands, 
add  that  the  crithidial  phase  takes  two  to  five  days. 

P.  Ill,  line  8  from  top  :  the  date  of  Sangiorgi  should  be  191 1. 

P.  142,  line  7  from  top  :  insert  "  Genus."  before  Eimeria. 

P.  252,  Insert  heading  "Family.  Opisthorchiidae,  Braun,  1901,"  above  "Sub-family. 
Opisthorchiinae,  Looss,  1899." 

P.  351,  description  of  fig.  255,  line  3  :  for  "  Thoma  "  read  "  Thomas." 

P.  471,  line  15  from  bottom  :  for  "  alcohol  100  parts  "  read  "alcohol  100  c.c. " 

P.  472,  line  II  from  bottom  :  for  "  Or  (2)  10  per  cent,  formalin^^^  read  "  Or  (2)  fix  in  hot 
10  per  cent,  formalin.''^ 

P.  493,  line  21  from  top  :  for  "  Conoy  "  read  "  Couvy." 

P.  589,  line  2  from  top  :  for  "  carnosa"  read  "  carnaria." 

P.  620,  line  15  from  top  :  for  "  fo  "  read  "  of." 

P.  622,  line  12  from  bottom  :  delete  comma  after  quantity. 

P.  626,  line  6  from  bottom  :  delete  comma  after  Mackie  (1915). 

P.  638  :  insert  title  "TREMATODES"  above  that  of  "  Fascioliasis. ' ' 

P.  709,  line  9  from  bottom  :  omit  second  Pediculus  capitis. 

P.  748,  line  8  from  top  :  for  "  cytologica  "  read  "  cy tological. " 

P.  753,  line  4  from  bottom  :  for  **  Fercocercous  "  read  **  Furcocercous." 

P.  755,  line  7  :  for  "  Oncocerca  "  read  "  Onchocerca." 


ON    PARASITES    IN    GENERAL, 


By  the  term  PARASITES  is  understood  living  organisms  which, 
for  the  purpose  of  procuring  food,  take  up  their  abode,  temporarily 
or  permanently,  on  or  within  other  Hving  organisms.  There  are 
both  phints  and  animals  (Phytoparasites  and  Zooparasites)  which 
lead  a  parasitic  life  in  or  upon  other  plants  and  other  animals. 

Phytoparasites  are  not  included  in  the  following  descriptions  of 
the  forms  of  parasitism,  but  a  very  large  number  of  animal  parasites 
(zooparasites)  are  described.  The  number  of  the  latter,  as  a  rule, 
is  very  much  underrated.  How  great  a  number  of  animal  parasites 
exists  may  be  gathered  from  the  fact  that  all  classes  of  animals  are 
subject  to  them.  Some  of  the  larger  groups,  such  as  Sporozoa,  Cestoda, 
Trematoda  and  Acaiithocephalaj  consist  entirely  of  parasitic  species, 
and  parasitism  even  occurs  among  the  vertebrates  {Myxine).  It 
therefore  follows  that  the  characteristics  of  parasites  lie,  not  hi  their 
structure,  but  in  the  manner  of  their  existence. 

Parasitism  itself  occurs  in  various  ways  and  degrees.  According 
to  R.  Leuckart,  we  should  distinguish  between  OCCASIONAL  (temporary) 
and  PERMANiiNT  (stationary)  PARASITISM.  Occasional  parasites,  such 
as  the  flea  {Pnlex  irritans),  the  bed-bug  {Cimex  lechUarius) ,  the  leech 
(Hinido  niedicinalis),  and  others,  only  seek  their  "  host  "  to  obtain 
nourishment  and  find  shelter  while  thus  occupied.  Without  being 
bound  to  the  host,  they  usually  abandon  the  latter  soon  after  the 
attainment  of  their  object  {Cimex,  Hirudo),  or  they  may  remain  on 
the  body  of  their  host  throughout  their  entire  development  from  the 
hatching  of  the  ^gg  {Pediculiis).  It  follows  from  this  mode  of  living 
that  the  occasional  parasites  become  sometimes  distinguishable  from 
their  free-living  relatives,  though  only  to  a  slight  extent.  It  is,  there- 
fore, seldom  difficult  to  determine  the  systematic  position  of  temporary 
parasites  from  their  structure. 

In  consequence  of  their  mode  of  life,  all  these  temporary  parasites 
live  on  the  external  surface  of  the  body  of  their  host,  though  more 
rarely  they  take  up  their  abode  in  cavities  easily  accessible  from  the 
exterior,  such  as  the  mouth,  nose  and  gills.  They  are  therefore 
frequently  called  Epizoa  or  Ectoparasites  ;  but  these  designations 


2  THE    ANJMAL   PARASITES   OF   MAN 

do  not  cover  only  the  temporary  parasites,  because  numerous  epizoa 
(as  for  instance  the  louse)  are  parasitic  during  their  entire  life. 

In  contradistinction  to  these  temporary  parasites,  the  permanent 
parasites  obtain  shelter  as  well  as  food  from  their  host  for  a  long 
period,  sometimes  during  the  entire  course  of  their  life.  They  do  not 
seek  their  host  only  when  requiring  nourishment,  but  always  remain 
with  it,  thus  acquiring  substantial  protection.  The  permanent  para- 
sites, as  a  rule,  live  within  the  internal  organs,  preferably  in  those 
which  are  easily  accessible  from  the  exterior,  such  as  the  intestine,  with 
its  appendages.  Nevertheless,  permanent  parasites  are  also  found  in 
separate  organs  and  systems,  such  as  the  muscular  and  vascular 
systems,  hollow  bones  and  brain,  while  some  live  on  the  outer  skin. 
Here  again,  the  terms  Entozoa  and  Endoparasites  do  not  include  all 
stationary  parasites  ;  to  the  latter,  for  instance,  the  lice  belong,  which 
pass  all  their  life  on  the  surface  of  the  body  of  their  host,  where  they 
find  shelter  and  food  and  go  through  their  entire  development.  The 
ectoparasitic  trematodes,  numerous  insects,  Crustacea,  and  other 
animals  live  in  the  same  manner. 

All  *^  Helminthes,"  however,  belong  to  the  group  of  permanent 
parasites.  This  term  is  now  applied  to  designate  certain  lowly  w^orms 
which  lead  a  parasitic  life  (intestinal  worms)  ;  but  they  are  not  all 
so  termed.  For  instance,  the  few  parasitic  Turbellaria  are  never 
classed  with  the  helminthes,  although  closely  related  to  them.  The 
turbellarians,  in  fact,  belong  to  a  group  of  animals  of  which  only 
a  few  members  are  parasitic,  whereas  the  helminthes  comprise  those 
groups  of  worms  of  which  all  species  (Cestoda,  Treinatoda,  Acantho- 
.cephala)f  or  at  least  the  majority  of  species  {Nematoda),  are  parasitic. 
Formerly  the  Linguatulidse  {Petitastoma)  were  classed  with  the 
helminthes  because  their  existence  is  also  endoparasitic,  and  because 
the  shape  of  their  body  exhibits  a  great  similarity  to  that  of  the  true 
helminthes.  vSince  the  study  of  the  development  of  the  Linguatulidae 
(P.  J.  van  Beneden,  1848,  and  R.  Leuckart,  1858)  has  demonstrated 
that  they  are  really  degenerate  arachnoids,  they  have  been  separated 
from  the  helminthes. 

It  is  hardly  necessary  to  emphasize  the  fact  that  the  helminthes 
or  intestinal  worms  do  not  represent  a  systematic  group  of  animals, 
but  only  a  biological  one,  and  that  the  helminthes  can  only  be  dis- 
cussed in  the  same  sense  as  land  and  water  animals  are  mentioned, 
i.e.y  without  conveying  the  idea  of  a  classification  in  such  a  grouping. 
It  is  true  that  formerly  this  was  universally  done,  but  very  soon  the 
error  of  such  a  classification  was  recognized.  Still,  until  the  middle 
of  last  century,  the  helminthes  were  regarded  as  a  systematic  group, 
although  C.  E.  V.  Baer  (1827)  and  F.  S.  Leuckart  (1827)  strenuously 
-opposed  this  view.     Under  the  active  leadership  of  J.  A.  E.  Goeze, 


PERMANENT   PARASITISM  3 

J.  G.  H.  Zeder,  J.  G.  Bremser,  K.  A.  Rudolphi  and  F.  Dujardin,  the 
knowledge  of  the  hehninthes  (hehninthology)  developed  into  a  special 
stud}',  but  unfortunately  it  lost  all  connection  with  zoology.  It 
required  the  intervention  of  Carl  Vogt  to  disestablish  the  helminthes 
as  one  class  of  animals,  by  uniting. the  various  groups  with  those  of 
the  free-living  animals  most  closely  related  to  them  {Platyhelminthes, 
N  ematlielinintlies). 

Permanent  parasitism  in  the  course  of  time  has  caused 
animals  adopting  this  mode  of  life  to  undergo  considerable,  some- 
times even  striking,  bodily  changes,  permanent  ectoparasites  having 
as  yet  undergone  least,  alteration.  The  latter  sometimes  bear  so 
unmistakably  the  likeness  to  the  group  to  which  they  belong,  that 
even  a  superficial  knowledge  of  their  structure  and  appearance  often 
suffices  for  the  recognition  of  their  systematic  position.  For  instance, 
though  the  louse,  like  many  decidedly  temporary  parasites,  has  lost 
its  wings — a  characteristic  of  insects — in  consequence  of  parasitism, 
yet  nobody  would  deny  its  insect  nature ;  such  also  occurs  in  other 
temporary  parasites  {Ciniex,  Pidex),  On  the  other  hand,  the  changes 
in  a  number  of  permanent  ectoparasites  (such  as  parasitic  Crustacea) 
are  far  more  considerable,  and  correspond  with  those  that  have 
occurred  in  permanent  endoparasites. 

These  alterations  depend  partly  on  retrogression  and  partly  on 
the  acquisition  of  new  peculiarities.  In  the  former  case,  the  change 
consists  in  the  loss  of  those  organs  which  have  become  useless  in  a 
permanent  parasitic  condition  of  existence,  such  as  wings  in  the 
louse,  and  the  articulated  extremities  seen  in  the  larval  stage  of 
parasitic  Crustacea.  The  loss  of  these  organs  goes  hand  in  hand 
with  the  cohesion  of  segments  of  the  body  that  were  originally 
separate,  and  alterations  in  the  muscular  and  nervous  systems.  In 
the  same  manner  another  means  of  locomotion  is  lost — the  ciliated 
coat — which  is  possessed  by  many  permanent  parasites  during  their 
larval  period.  To  all  appearances,  this  character  is  not  secondary 
and  recently  acquired,  but  represents  a  primary  character  inherited 
from  free-living  progenitors,  and  still  transmitted  to  the  altered 
descendants,  because  of  its  use  during  the  larval  stage  (e.g.,  the  larvae 
of  a  great  many  Trematodes,  the  oncospheres  of  some  Cestodes). 
Amongst  the  retrogressions,  the  loss  of  the  organs  of  sense  may 
be  mentioned,  particularly  the  eyes,  which  are  still  present,  not  only 
in  the  nearest  free-living  forms  but  also  in  the  free-living  larvae  of 
true  parasites.  It  is  only  quite  exceptionally  that  the  eyes  are 
subsequently  retained,  as  a  rule  they  are  lost.  Lastly,  in  a  great 
many  cases  the  digestive  system  also  disappears,  as  in  parasitic 
Crustacea,  in  a  few  nematodes  and  trematodes,  in  all  cestodes  and 
Acanthocephala.  There  remain  at  most  the  rudiments  of  the  muscles 
of  the  fore -gut,  but  these  are  adapted  to  entirely  different  uses. 


4  THE   ANIMAL   PARASITES   OF   MAN 

The  new  characters  which  permanent  parasites  may  acquire  are, 
first  of  all,  the  remarkably  manifold  clasping  and  clinging  organs, 
which  are  seldom  (as  in  parasitic  Crustacea)  directly  joined  on  to 
already  existing  structures.  In  those  instances  in  which  organs 
for  the  conveyance  of  food  are  retained,  these  likewise  frequently 
undergo  transformation,  in  consequence  of  the  altered  food  and 
manner  of  feeding.  Such  alterations  consist,  for  instance,  in  the 
transformation  of  a  masticating  mouth  apparatus  into  the  piercing 
and  sucking  organs  of  parasitic  insects. 

Hermaphroditism  (as  m  Trematodes,  Cestodes,  and  a  few  Nema- 
todes) is  a  further  peculiarity  of  many  permanent  parasites  ;  moreover, 
the  association  in  couples  that  occurs,  especially  in  trematodes,  may 
lead  to  complete  cohesion  and,  exceptionally,  also  to  re-separation 
of  the  sexes.  In  many  cases  the  females  only  are  parasitic,  while 
the  males  live  a  free  life,  or  there  may  be  in  addition  the  so-called 
complementary  males.  Occasionally  the  male  alone  is  parasitic,  and 
in  that  case  lives  within  the  female  of  the  same  species,  which  may 
live  free,  like  certain  Gephyrea  {Bonellia)  ;  or  the  female  also  may 
be  parasitic,  as  Trichosoma  crassicaudum,  which  lives  in  the  bladder 
of  the  sewer  rat  {Miis  decumanns). 

We  have  numerous  proofs  that  demonstrate  how  considerably 
the  original  features  of  many  parasites  have  become  changed.  We 
need  only  draw  attention  to  the  aforementioned  Linguatulidae,  also 
to  many  of  the  parasitic  Crustacea  belonging  to  various  orders.  In 
all  of  these  a  knowledge  of  the  larval  stages — in  which  there  is  no 
alteration,  or  at  most  only  a  slight  degree  of  change — serves  to 
determine  their  systematic  position,  i.e.^  the  nearest  conditions  of 
relationship. 

The  most  remarkable  changes  are  observed  in  those  groups  that 
contain  only  a  few  parasitic  members,  the  majority  leading  a  free 
life.  A  striking  instance  is  afforded  by  a  snail,  the  well-known  Ento- 
concha  mirabilts,  Miiller.  This  mollusc  consists  merely  of  an  elongated 
sac  living  in  a  Holothurian  {Synapta  digitata).  It  possesses  none 
of  the  characteristics  of  either  the  Gastropoda  or  any  molluscs, 
and  in  its  interior  there  is  nothing  to  be  observed  but  the  organs 
of  generation  and  the  embryos.  Nevertheless,  the  Entoconcha  is 
decidedly  a  parasitic  snail,  as  is  clearly  proved  by  its  larvae,  but 
it  is  a  snail  which,  in  consequence  of  parasitism,  has  lost  all  the 
characteristics  of  molluscs  in  its  mature  condition,  but  still  exhibits 
them  in  the  early  stages  of  development. 

Certain  nematodes  show  very  clearly  to  what  devious  courses 
parasitism  may  lead.  The  Atractonema  gibbosnin,  the  life-history  of 
which  has  been  described  by  R.  Leuckart,  and  which  lives  in  the 
larvae  and  pupae  of   a  dipterous  insect  (Cecidoniyia),  exhibits,   in  its 


FERTILITY   OF   PARASITES  5 

early  stage,  the  ordinary  characteristics  of  other  threadworms.  A 
few  weeks  later — the  males  having  died  off  immediately  after  copu- 
lation— the  females  are  transformed  into  spindle-shaped  bodies,  the 
mouth  and  anus  of  which  are  closed.  They  carry  with  them  an  irregu- 
larly shaped  appendage,  in  which  the  segmenting  ova  are  situated, 
and  in  which  the  further  conditions  of  life  of  the  Atradotiema  are 
accomplished.  A  minute  examination  has  demonstrated  that  this 
appendage  is  the  prolapsed  and  enlarged  vagina  of  the  animal  which 
has  become  merely  a  supplementary  attachment.  The  conditions 
present  in  the  Sphcendaria,  the  nematoid  nature  of  which  was  long 
undiscovered,  are  still  more  remarkable.  It  was  only  when  Siebold 
proved  that  typical  nematodes  were  hatched  from  their  eggs 
that  their  nature  was  recognized.  The  nematodes  thus  produced 
have  not  the  slightest  resemblance  to  the  parent. 

The  researches  of  Lubbock,  A.  Schneider,  and  more  particularly 
of  R.  Leuckart,  have  shown  that  what  we  call  Sphcerularia  homhi  is  not 
an  animal  but  merely  an  organ — the  vagina — of  a  nematode  worm. 
This  vagina  at  first  grows,  sac-like,  from  the  body  of  the  tiny  nema- 
tode ;  it  gradually  assumes  enormous  dimensions  (2  cm.  in  length)  ; 
it  contains  the  sexual  organs  and  parts  of  the  intestine.  The  remain- 
ing portion  of  the  actual  animal  then  becomes  small  and  shrivelled  ; 
it  may  be  easily  overlooked,  being  but  an  appendage  to  the  vagina 
with  its  independent  existence,  and  it  finally  disappears  altogether. 

The  GREAT  FERTILITY  of  parasites  is  another  of  their  peculiarities, 
though  this  may  be  also  the  case  to  a  certain  degree  with  some  of 
the  free-living  animals,  the  progeny  of  which  are  likewise  exposed 
to  enormous  destruction. 

More  remarkable,  however,  is  the  fact  that  the  young  of  the 
endoparasites  only  very  exceptionally  grow  to  maturity  by  the  side 
of  their  parents.  Sooner  or  later  they  leave  the  organ  inhabited  by 
tlie  parents,  frequently  reach  the  open,  and  after  a  shorter  or  longer 
period  of  free  existence  seek  new  hosts.  During  their  free  period, 
moreover,  a  considerable  growth  may  be  attained,  or  metamorphosis 
may  take  place,  or  even  multiplication.  In  the  exceptional  cases  in 
which  the  young  remain  within  the  same  host,  they  nevertheless 
usually  quit  the  organ  inhabited  by  the  parents.  They  likewise  rarely 
attain  maturity  within  the  host  inhabited  by  the  parents,  but  only, 
as  in  other  cases,  after  having  gained  access  to  fresh  hosts. 

These  transmigrations  play  a  very  important  rdle  in  the  natural 
history  of  the  internal  parasites,  but  they  frequently  conceal  the  cycle 
of  development,  for  sometimes  there  are  INTERMEDIATE  GENERATIONS, 
which  themselves  invade  intermediate  hosts.  Even  when  there  are  no 
intermediate  generations,  the  system  OF  intermediate  HOSTS  is 
frequently  maintained  by  the  endoparasites. 


6  THE   ANIMAL   PARASITES   OK   MAN 

According  to  the  kind  of  food  ingested  by  parasites,  it  has 
recently  become  usual  to  separate  the  true  parasites  from  those 
animals  that  feed  on  the  superfluity  of  the  food  of  the  host,  or  on 
products  which  are  no  longer  necessary  to  him,  and  to  call  the  latter 
MESSMATES  or  COMMENSALS.  As  examples,  the  Ricinidae  are  thus 
designated,  because,  like  actual  lice,  they  dwell  among  the  fur  of 
mammals  or  the  plumage  of  birds.  They  do  not,  however,  suck 
blood,  for  which  their  mouth  apparatus  is  unsuited,  but  subsist 
on  useless  epidermic  scales.  These  epizoa,  according  to  J.  P.  van 
Beneden,  are,  to  a  certain  extent,  useful  to  their  hosts  by  removing 
deciduous  materials  which  under  certain  circumstances  might  become 
harmful  to  them.^  This  investigator,  who  has  contributed  so  greatly 
to  our  knowledge  of  parasites,  assigns  the  Ricines  to  the  MUTUALISTS, 
under  which  term  he  comprises  animals  of  various  species  which 
live  in  common,  and  confer  certain  benefits  on  one  another.  The 
mutualists  are  usually  intimately  connected  in  a  mutually  advan- 
tageous association  known  as  '^  symbiosis."^ 

Incidental  and  Pseudo  Parasites. — In  many  cases  the  parasites  are 
confined  to  certain  hosts,  and  may  therefore  be  designated  as  specific 
to  such  hosts.  Thus,  hitherto,  Tcenia  solinm  and  Tcenia  saginata 
in  their  adult  condition  have  only  been  found  in  man  ;  Tcenia  crassi- 
collis  only  in  the  cat ;  Brandesia  (Distoma)  turgida  and  Halipegus 
{Distoma)  ovocaudatus  only  in  Rana  esculenta,  and  so  forth.  In  many 
other  cases,  however,  certain  species  of  parasites  are  common  to  several, 
and  sometimes  many,  species  of  hosts  ;  Dipylidium  caninnm  is  found 
in  the  domestic  cat  as  well  as  in  the  dog;  Fasciola  hepatica  is  found  in 
a  large  number  of  herbivorous  mammals  (nineteen  species),  Diplodiscus 
(Amphistomunt)  suhclavatus  in  numerous  urodele  and  ecaudate  am- 
phibia, Holostomnm  variahile  in  about  twenty-four  species  of  birds, 
and  so  on.  In  these  cases  the  hosts  are  almost  invariably  closely  related, 
belonging,  as  a  rule,  to  the  same  family  or  order,  or  at  any  rate  to  the 
same  class.  Trichinella  spiralis^  which  is  found  in  man,  and  in  the  pig, 
bear,  rat,  mouse,  cat,  fox,  badger,  polecat  and  marten,  and  is  capable 
of  being  artificially  cultivated  in  the  dog,  rabbit,  sheep,  horse,  in 
other  mammals,  and  even  in  birds,  is  one  of  the  most  striking 
exceptions. 

Some  parasites  are  so  strictly  confined  to  one  species  of  host  that, 
even  when   artificially  introduced  into   animals   very  closely  related 


^  According  to  Sambon,  the  Ricinidae  are  by  no  means  advantageous  to  their  hosts.  These 
Hemipterous  parasites  give  rise  to  an  intolerable  itching  which  may  cause  loss  of  rest,  emacia- 
tion, and  sometimes  even  death.  Birds  suffering  from  phthiriasis  of  the  Ricines  are  usually 
in  bad  health. 

-  For  further  information  on  these  conditions,  see  "Die  Schmarotzer  des  Thierreichs,"  by 
P.  J.  van  Beneden,  Leipzig,  1876  ;  and  *'  Die  Symbiose,"  by  O.  Hertwig. 


INCIDENTAL   AND   PSEUDO-PARASITES  7 

to  their  normal  host,  they  do  not  thrive,  but  sooner  or  later,  often 
very  quickly,  die  off,  and  very  rarely  establish  themselves.  For 
example,  repeated  attempts  have  been  made  to  rear  the  adult  Tcenia 
soli  mil  in  the  dog,  or  to  rear  Cysticercus  cellulosce  in  the  ox,  or  the 
Cyst icerc lis  of  Tcenia  saginata  in  the  pig,  but  they  have  always  proved 
unsuccessful.  Only  exceptionally  has  it  been  possible  to  transfer 
Ccenunis  cerebralis,  the  larval  stage  of  a  tapeworm  (Tcenia  ccenurus)  of 
the  dog  from  the  brain  of  the  sheep  to  that  of  the  domestic  goat. 
On  the  other  hand,  in  the  case  of  the  Trichinellae  transference  to 
a  different  host  is  easily  accomplished. 

Under  natural  conditions,  it  is  not  uncommon  for  certain  kinds 
of  specific  parasites  to  occur  occasionally  in  unusual  hosts.  Their 
relationship  to  the  latter  is  that  of  incidental  parasites.  Thus 
Echinorhynchiis  gigas,  a  specific  parasite  of  the  pig,  is  only  an 
incidental  parasite  of  man  ;  Fasciola  hepatica  and  Dicroccelium 
lanceatnm  are  specific  to  numerous  kinds  of  mammals,  but  may 
be  found  incidentally  in  man.  On  the  other  hand,  Dibothriocephalus 
latuSj  a  specific  parasite  of  man,  may  occasionally  take  up  its  abode 
in  the  dog,  cat  and  fox.  As  a  rule,  all  those  parasites  of  man  that 
are  only  rarely  met  with,  notwithstanding  that  human  beings  are 
constantly  being  observed  and  examined  by  medical  men,  are 
termed  incidental  parasites  of  man.  In  many  cases  we  are 
acquainted  with  the  normal  or  specific  host  of  these  parasites. 
Thus  we  know  the  specific  host  of  Balantidium  colt,  Eimeria  stiedcBf 
Fasciola  hepatica,  Dipylidiuin  caninuin,  etc. ;  in  others  the  host  is  as 
yet  unknown.  In  the  latter  case  the  question  partly  relates  to  such 
forms  as  have  been  so  deficiently  described  that  their  recognition 
is  impossible,  partly  to  parasites  of  man  in  various  regions  of  the 
earth,  the  Helminthes  and  parasites  of  which  are  totally  unknown 
or  only  slightly  known,  or  finally  to  early  developmental  stages  that 
are  difficult  to  identify.  Animals  that  usually  live  free,  and  exception- 
ally become  parasitic,  may  likewise  be  called  incidental  parasites. 
In  this  category  are  included  a  few  Angiiillulidce  that  have  been 
observed  in  man  ;  also  Leptodera  appendicidata,  which  usually  lives 
free,  but  may  occasionrJly  become  parasitic  in  black  slugs  {Arion 
einpiricoriun)  :  when  parasitic  it  attains  a  larger  size,  and  produces 
far  more  eggs  than  when  living  a  free  life.  In  order  to  avoid  errors, 
the  term  "  incidental  parasites  "  should  be  confined  to  true  parasites 
which,  besides  living  in  their  normal  host,  may  also  live  in  other 
hosts.  Leuckart  speaks  of  facultative  parasitism  in  such  forms 
as  Leptodera.  L.  Oerley^  succeeded  in  artificially  causing  Leptodera 
(Rhabditis)  pellio  to  assume  facultative  parasitism  by  introducing  these 

*  Oerley,  L.,  *'  Der  Rhabditiden  und  ihre  medizinische  Bedeutung,"  Berlin,  1886,  p.  65. 


8  THE   ANIMAL   PARASITEvS   OF  MAN 

worms  into  the  vagina  of  mice,  where  the  parasites  remained  aHve  and 
multipHed.  Lcptodera  pellio  dies  in  the  intestines  of  mammals  and 
man  ;  it  remains  alive  in  frogs,  but  always  escapes  into  the  open 
with  the  faeces. 

Recently  the  incidental  parasites  of  man  have  also  been  called 
"Pseudo-parasites"  or  "  Pseudo-helminthes."  Formerly,  how- 
ever, these  terms  were  applied  not  only  to  living  organisms  that  do 
not  and  cannot  live  parasitically,  and  that  only  exceptionally  and 
incidentally  get  into  man,  but  also  to  any  foreign  bodies,  portions 
of  animals  and  plants,  or  even  pathological  formations  that  left  the 
human  system  through  the  natural  channels,  and  the  true  nature 
of  which  was  misunderstood.  Frequently  these  bodies  were  described 
as  living  or  dead  parasites  and  labelled  with  scientific  names,  as  if 
they  w^ere  true  parasites.  A  study  of  these  errors,  which  formerly 
occurred  very  frequently,  w^ould  be  as  interesting  as  it  would  be 
instructive.  It  is  better  not  to  use  the  expression  pseudo-parasites  for 
incidental  parasites,  but  to  keep  to  the  original  meaning,  for  it  is  not 
at  all  certain  that  pseudo-parasites  are  not  described,  even  nowadays. 

The  Influence  of  Parasites  on  the  Host. — In  a  great  many  cases, 
we  are  not  in  a  position  to  state  anything  regarding  any  marked 
influence  exercised  by  the  parasite  on  the  organism,  and  on  the  con- 
ditions of  life,  of  the  host.  Most  animals  and  many  persons  exhibit 
few  signs  of  such  influence,  an  exception  being  infestation  with 
helminthes  and  certain  other  parasites  w^hich  produce  eosinophilia 
in  the  blood.  As  a  general  rule,  the  parasite,  which  is  always  smaller 
and  weaker  than  its  host,  does  not  attempt  to  endanger  the  life 
of  the  latter,  as  simultaneously  its  own  existence  would  be  threatened. 
The  parasite,  of  course,  robs  its  host,  but  usually  in  a  scanty  and 
sparing  manner,  and  the  injuries  it  inflicts  can  hardly  be  taken  into 
account.  There  are,  however,  numerous  cases^  in  which  the  situation 
of  the  parasites  or  the  nature  of  their  food,  added  to  their  number  and 
movements,  may  cause  more  or  less  injury,  and  even  threaten  the  life 
of  the  host.  It  stands  to  reason  that  a  Cysticercus  cellulosce  situated  in 
the  skin  is  of  but  slight  importance,  whereas  one  that  has  penetrated 
the  eye  or  the  brain  must  give  rise  to  serious  disorders.  A  cuticular 
or  intestinal  parasite  is,  as  a  rule,  less  harmful  than  a  blood  parasite. 
A  helminth,  such  as  an  Ascaris  lunibricoides  or  a  tapeworm,  that 
feeds  on   the  residues  of  foodstuffs  w^ithin  the  intestine,  will  hardly 


'  Liihe,  M.,  "  Ueber  d.  Fix.  d.  Helm.  a.  d.  Darmwand  ihrer  Wirthe  u.  die  dadurch 
verursachten  path-anat.  Veranderungen  d.  Wirthsdarmes,"  Trans,  of  IVth  Intern.  Zool. 
Cong.,  Berlin,  1901  ;  Mingazzini,  P.,  '*  Ric.  sul  var.  modo  di  fiss.  delle  tenie  alia  par.  int. 
e  sul  loro  assorbimento,"  Ric.  Lab.  Anat.  Roma  e  altri  Lab.  bioL,  vol.  x,  1904;  Shipley, 
A.  E.,  and  E.  G.  Fearnsides,  "The  Effects  of  Metazoan  Parasites  on  their  Hosts,"  /ottrn, 
Econ.  Biol.,  1906,  i,  2. 


THE   INFLUENCE   OF   PARASITES  9 

afifect  its  host  by  depriving  it  of  this  material.  The  case  is  different 
when  the  parasites  are  very  numerous,  especially  when  the  heavily 
infested  host  happens  to  be  a  young  individual  needing  all  it  ingests 
for  its  own  requirements,  and  therefore  unable  to  sustain  the  drain  of 
numerous  intruders  in  the  intestine.  Disturbances  also  set  in  more 
rapidly  when  the  intestinal  helminthes  are  blood-suckers,  the  injury 
to  the  host  resulting  from  the  kind  of  food  taken  by  the  parasite. 

Generally,  the  disorders  caused  by  loss  of  chyle  are  insignificant 
when  compared  with  those  induced  by  the  growth  and  agglomera- 
tion of  the  helminthes.  The  latter  may  cause  chiefly  obstructions  of 
small  vessels  or  symptoms  of  pressure  in  affected  or  contiguous  organs, 
with  all  those  complications  which  may  arise  secondarily,  or  they  may 
even  lead  to  the  complete  obliteration  of  the  organ  invaded.  Of  course 
the  symptoms  will  vary  according  to  the  nature  of  the  organ  attacked. 

In  consequence  also  of  the  movements  of  the  parasites,  disorders 
are  set  up  that  may  tend  to  serious  pathological  changes  of  the 
affected  organs.  The  collective  migrations,  undertaken  chiefly  by 
the  embryos  of  certain  parasites  (as  in  trichinosis,  acute  cestode 
tuberculosis),  are  still  more  harmful,  as  are  also  the  unusual  migrations 
•of  other  parasites,  which,  incidentally,  may  lead  to  the  formation 
of  so-called  worm  abscesses  or  to  abnormal  communications  (fistulae) 
between  organs  that  are  contiguous  but  possess  no  direct  connection. 

Recently,  several  authors  have  called  attention  to  the  fact  that 
the  helminthes  produce  substances  that  are  toxic  to  their  host ;  and 
II le  effects  of  such  poisons  explain  the  pathology  of  helminthiasis 
far  more  satisfactorily  than  the  theory  of  reflex  action. 

In  a  number  of  cases  these  toxic  materials  (leucomaines)  have  been 
isolated  and  their  effects  on  living  organisms  demonstrated  by  actual 
experiments.  It  also  appears  that  the  absorption  of  materials  formed 
by  the  decomposition  of  dead  helminthes  may  likewise  cause  toxic 
effects.  However,  our  knowledge  of  these  conditions  is  as  yet  in 
its  initial  stage. ^ 

Nearly  all  the  symptoms  caused  directly  or  indirectly  by  parasites 
are  of  such  a  nature  that  the  presence  of  the  parasites  cannot  be 
diagnosed  with  any  certainty,  or  only  very  rarely.  The  most  that 
can  be  done  is  to  deduce  the  presence  of  parasites  by  the  exclusion 
of  other  causes.     Fortunately,   however,  there   are  sufficient   means 


1  Moursson  et  Schlagdenhaufifen,  "Nouv.  rech.  clin.  et  phys.  sur  quelq.  liquides  organ.,*' 
€.  R.  Acad.  Set.,  Paris,  1882,  p.  791  ;  Debove,  '*  Del'intox.  hydat.,"^«//.  et  Mint.  Soc.  m'ed.  des 
Hopit.,  1888  ;  Linstow,  v.,  "  Ueb.  d.  Giftgehalt  d.  helm.,"  Internat.  Monatsschr.  f.  Anat.  u. 
Phys.,  xiii,  1896;  Peiper,  **Z.  Symptomatol.  der  thier.  Paras.,"  Deutsche  med.  Wochenschr., 
1897,  No.  40;  Mingazzini,  P.,  *' Ric.  sul  veleno  d.  elm.  int.,"  Rass.  intern,  d.  med. 
modern.  Ann.,  1901,  ii,  No.  6;  Vaullegeard,  A.,  "  Etud.  exp.  et  crit.  sur  Taction  d.  helm.," 
Btill.  Soc.  Linn,  de  Normandie,  1901,  5,  Ser.  T,  vii,  p.  84,  and  others. 


lO  THE   ANIMAL   PARASITES   OF   MAN 

by  which  we  may  confirm  the  diagnosis  in  a  great  many  cases.  Such 
means  consist  not  only  in  a  minute  examination  of  the  patient  by 
palpation,  percussion  and  local  inspection,  but  also  in  the  micro- 
scopical examination  of  the  natural  secretions  and  excretions  of  the 
body,  such  as  sputum,  nasal  mucus,  urine  and  fpeces.  Though  such 
examinations  may  entail  loss  of  time,  they  are  necessary  in  the  interest 
of  the  patient.  It  appears,  moreover,  that  quackery,  which  has 
gained  considerable  ground  even  in  the  treatment  of  the  helminthic 
diseases  of  man,  can  thus  be  considerably  limited. 

Origin  of  Parasites} — In  former  times,  when  the  only  correct  views 
that  existed  related  to  the  origin  of  the  higher  animals,  the  mode 
of  multiplication  of  parasites  as  well  as  of  other  lowly  animals 
was  ascribed  to  spontaneous  generation  {geueratio  ceqnivoca),  and 
this  opinion  prevailed  throughout  the  middle  ages.  The  writers  on 
natural  science  merely  devoted  their  time  to  the  interpretation  of  the 
views  of  the  old  authors,  and  perpetuated  the  opinions  of  the  ancients 
on  questions,  which,  even  in  those  days,  could  have  been  correctly 
explained  merely  by  observation. 

It  was  only  w^hen  observations  were  again  recommenced,  and  the 
microscope  was  invented,  that  the  idea  of  spontaneous  generation 
became  limited.  Not  only  did  the  microscope  reveal  the  organs  of 
generation  or  their  products  (eggs)  in  numerous  animals,  but  Redi 
succeeded  in  proving  that  the  so-called  Helcophagi  (flesh  maggots)  are 
only  the  progeny  of  flies,  and  never  appear  in  the  flesh  of  slaughtered 
animals  when  fully  developed  flies  are  prevented  from  approaching 
and  depositing  their  eggs  on  it.  Swammerdam  likewise  knew  that 
the  *^ worms"  living  in  the  caterpillars  of  butterflies  were  the  larvae 
of  other  insects  (ichneumon  flies)  which  had  laid  their  eggs  in  their 
bodies  ;  he  also  discovered  the  ova  of  lice.  The  tw^o  authors  men- 
tioned were,  however,  unwilling  to  see  that  the  experience  they  had 
gained  regarding  insects  applied  to  the  helminthes.  Leeuwenhoek 
also  vehemently  opposed  the  theory  of  a  spontaneous  generation, 
maintaining  that,  on  a  basis  of  common-sense,  eggs,  or  at  all  events 
germs,  must  exist,  even  though  they  could  not  be  seen. 

The  use  of  the  microscope  also  revealed  a  large  number  of  very 
small  organisms  in  the  water  and  moist  soil,  some  of  which  un- 
doubtedly resembled  helminthes.  Considering  the  wide  dissemination 
of  these  minute  organisms,  it  was  natural  to  conjecture  that  after  their 
almost  unavoidable  introduction  into  the  human  system  they  should 
grow  into  helminthes  (Boerhave,  Hoffmann).  Linnaeus  went  even 
further,  for  he  traced   the  descent  of  the  liver-fluke  of  sheep  from 

'  Die  Geschichte  der  *'  Klinisch  wichtigen  Parasiten,"  behandelt  H.  Vierordt  im  **Handb. 
d.  Gesch.  d.  Med.  hrsg."  v.  M.  Neuburger  u.  J.  Pagel,  Bd.  ii,  1903. 


ORIGIN   OF   PARASITES  II 

a  free-living  planaria  {Dendroccelum  lacteuui),  the  Oxyuris  verniicnlaris 
from  free-living  nematodes,  and  the  Tcenia  lata  (i.e.,  Dibothriocephalus 
latus)  from  a  tapeworm  {Schistocephalits  solidns)  found  free  in  the 
water.  Linnaeus'  statements  met  with  general  approval.  However, 
we  must  bear  in  mind  that  at  that  time  the  number  of  helminthes 
known  was  very  small,  and  many  of  the  forms  that  we  have  long 
ago  learned  to  differentiate  as  specific  were  then  regarded  as  belonging 
to  one  species.  Linnaeus'  statements  were  partly  supported  by  similar 
discoveries  by  other  investigators,  such  as  Unzer,  and  partly  also  by 
the  discovery  of  eggs  in  many  helminthes.  It  was  believed  that  the 
eggs  hatched  in  the  outside  world  gave  rise  to  free-living  creatures, 
and  that  these,  after  their  introduction  into  the  intestine,  were  trans- 
formed into  helminthes.  By  means  of  these  eggs  the  old  investigators 
tried  to  explain  the  hereditary  transmission  of  the  intestinal 
worms,  which  was  universally  believed  until  the  commencement  of 
the  last  century.  Some  authors  went  so  far  as  to  regard  the  intestinal 
worms  as  congenital  or  inherited  ;  they  maintained  the  possibility 
of  direct  transmission,  as  in  suckling,  and  denied  that  the  eggs 
reaching  the  external  world  had  anything  to  do  with  the  propagation 
of  the  parasites. 

The  more  minute  comparison  between  the  supposed  free-living 
stages  of  the  helminthes  and  their  adult  forms,  and  the  impossibility 
of  finding  corresponding  free  forms  for  the  ever-increasing  number 
of  parasitic  species,  revealed  the  improbability  of  Linnaeus'  statements 
(O.  Fr.  Miiller).  It  was  the  latter  author  also  who  recognized  the 
origin  of  the  tapeworms  {Schistocephalits,  Ligtila)  found  free  in  the 
water.     They  originate  from  fishes  which  they  quit  spontaneously. 

However,  in  spite  of  the  fact  that  van  Doeveren  and  Pallas 
correctly  recognized  the  significance  of  the  eggs  in  the  trans- 
mission of  intestinal  worms,  these  statements  remained  disregarded, 
as  did  Abildgaard's  observation,  experimentally  confirmed,  that  the 
(immature)  cestodes  from  the  abdominal  cavity  of  sticklebacks  became 
mature  in  the  intestines  of  aquatic  birds.  Moreover,  at  the  end  of 
the  eighteenth  and  the  commencement  of  the  nineteenth  centuries, 
after  helminthology  had  been  raised  to  a  special  branch  of  study 
by  the  successful  results  of  the  investigations  of  numerous  authors 
(Goeze,  Bloch,  Pallas,  Miiller,  Batsch,  Rudolphi,  Bremser),  many 
of  whom  experienced  a  ^'  divine  joy  "  in  searching  the  intestines  of 
animals  for  helminthes,  some  authors  reverted  to  generatio  a^quivoca, 
without,  however,  entirely  denying  the  existence  of  organs  of  genera- 
tion and  eggs.  The  fact  that  a  few  nematodes  bore  living  progeny — 
a  fact  of  which  Goeze  w^as  already  aware — had  no  influence  on  the 
erroneous  opinion,  as  in  such  cases  it  was  considered  that  the  young 
continued    to    develop    beside    the    old    forms.      There    were    also 


12  THE   ANIMAL   PARASITES   OF   MAN 

many  helminthes  known  that  never  developed  sexual  organs  and 
never  produced  eggs,  and  which  therefore  were  referred  to  generatio 
cequivoca.  People  were  convinced  that  the  intestinal  mucous  mem- 
brane or  an  intestinal  villus  could  transform  itself  into  a  worm, 
either  in  a  general  morbid  condition  of  the  body,  or  in  pathological 
changes  of  a  more  local  character.  The  appearance  of  helminthes 
was  even  regarded  as  useful  and  as  a  means  for  the  expulsion  of 
injurious  matter. 

These  views,  firmly  rooted  and  supported  by  such  eminent 
authorities  as  Rudolphi  and  Bremser,  could  not  easily  be  overthrown. 
First,  a  change  took  place  in  the  knowledge  of  the  trematodes. 
In  1773,  O.  Fr.  Miiller  discovered  Cercaricv  living  free  in  water. 
He  regarded  them  as  independent  creatures  and  gave  them  the  name 
that  is  still  used  at  the  present  time.  Nitzsch,  who  also  minutely 
studied  these  organisms  and  who  recognized  the  resemblance  of 
the  anterior  part  of  their  bodies  to  a  Fasciola,  did  not,  however,  arrive 
at  a  correct  conclusion.  He  regarded  the  combination  rather  as 
that  of  a  Fasciola  with  a  Vibrio,  for  which  he  mistook  the  charac- 
teristic tail  of  the  cercaria.  He  also  noticed  the  encystment  (trans- 
formation into  the  *'pupa")  on  foreign  bodies  of  many  species  of 
these  animals,  but  was  of  opinion  that  this  process  signified  only  the 
termination  of  Hfe. 

Considerable  attention  was  attracted  to  the  matter  when  Bojanus 
first  published  a  paper  entitled  ''A  Short  Note  on  Cercaria  and  their 
Place  of  Origin."  He  pointed  out  that  the  cercariae  creep  out  of  the 
''royal  yellow^  worms,"  which  occur  in  freshwater  snails  {LuiuKxa, 
Pakidina)y  and  are  probably  generated  in  these  worms. 

Oken,  in  whose  journal,  his  (1818,  p.  729),  Bojanus  published  his 
discovery,  remarks  in  an  annotation,  "  One  might  lay  a  wager  that 
these  Cercariae  are  the  embryos  of  Distomes."  Soon  after  (1827), 
C.  E.  V.  Baer  was  able  to  confirm  Bojanus'  hypothesis  that  the 
cercariae  as  a  ''  heterogeneous  brood  "  originated  from  spores  in  para- 
sitic tubes  in  snails  (germinating  tubes).  Moreover,  Mehlis  [his,  183 1, 
p.  190)  not  only  discovered  the  opercula  of  the  ova  of  Distoma,  but 
likewise  saw  the  infusorian-like  embryo  emerge  from  the  eggs  of 
Typhloccelmn  {Monostomum)  flaviim  and  Cathcemasia  {Distoma)  hians. 
A  few  years  later  (1835)  v.  Siebold  observed  the  embryos  (miracidia)  of 
the  Cyclocoelnm  {Monostomum)  mutabile,  and  discovered  in  their  interior 
a  cylindrical  body  that  behaved  like  an  independent  being  ("  necessary 
parasite"),  and  was  so  similar  in  appearance  to  the  ''royal  yellow 
worms  "  (Bojanus)  that  Siebold  considered  the  origin  of  the  latter  from 
the  embryos  of  trematodes  as,  at  all  events,  possible.  Meanwhile, 
V.  Nordmann  of  Helsingfors  had  in  1832  seen  the  miracidia  of  flukes 
provided  with  eyes  swimming  in  water;  v.  Siebold  (1835)  had  observed 


ORIGIN    OF   PARASITES  13 

the  embryos,  or  oncospheres,  of  tapeworms  furnished  with  six 
booklets  in  the  so-called  eggs  of  the  Taenia;- while  Creplin  (1837)  had 
discovered  the  'Mnfusorial"  young  of  the  Diphyllobothrium  {Bothrio- 
cephahis)  ditrcninui,  and  conjectured  that  similar  embryos  were  to  be 
found  in  other  cestodes  with  operculated  eggs.  At  all  events,  the  fact 
was  established  that  the  progeny  of  the  helminthes  appeared  in  various 
forms  and  was  partly  free  living.  The  researches  of  Eschricht  (1841) 
were  likewise  of  influence,  as  they  elucidated  the  structure  of  the 
Bothriocephali,  and  proved  that  the  encysted  and  sexless  helminthes 
were  merely  immature  stages. 

J.  I.  Steenstrup  (1842)  was,  however,  the  first  to  furnish  explana- 
tions for  the  numerous  isolated  and  uncomprehended  discoveries. 
Commencing  with  the  remarkable  development  of  the  Coelenterata, 
he  established  the  fact  that  the  Helminthes,  especially  the  endopara- 
sitic  trematodes,  multiply  by  means  of  alternating  and  differently 
formed  generations.  Just  as  the  polyp  originating  from  the  Qgg  of 
a  medusa  represents  a  generation  of  medusae,  so  does  the  germinal 
tube  ('^  royal  yellow  w^orm  ")  originating  from  the  ciliated  embryo  of 
a  Distoma,  etc.,  represent  the  cercaria.  These  w^ere  consequently 
regarded  as  the  progeny  of  trematodes,  and  Steenstrup,  guided  by 
his  observations,  conjectured  that  the  cercaria,  whose  entrance  into 
the  snails  he  had  observed  accompanied  by  the  simultaneous  loss  of 
the  propelling  tail,  finally  penetrated  into  other  animals,  in  which  they 
became  flukes. 

Part  of  this  hypothetical  cycle  of  development  was  erroneous,  and 
in  other  particulars  positive  observation  was  lacking,  but  the  path 
pursued  was  in  the  right  direction.  Immediately  after  the  appear- 
ance of  Steenstrup's  celebrated  work,  v.  Siebold  expressed  his  opinion 
that  the  encapsuled  flukes  certainly  had  to  travel,  i.e.y  to  be  trans- 
mitted with  their  bearers  into  other  hosts,  before  becoming  mature. 
This  view  was  experimentally  confirmed  by  de  Filippi,  La  Valette 
St.  George  (1855),  as  w^ell  as  by  Pagenstecher  (1857),  while  the  meta- 
morphosis of  the  ciliated  embryo  of  Distoma  into  a  germinal  tube 
was  first  seen  by  G.  Wagener  (1857)  in  Gorgodera  (Distoma)  cygnoides 
of  frogs.  All  that  we  have  subsequently  learned  from  the  works  of 
numerous  investigators  about  the  development  of  endoparasitic  trema- 
todes has  certainly  increased  our  knowledge  in  various  directions,  and, 
apart  from  the  deviating  development  of  the  Holostomidce  has,  as  a 
whole,  confirmed  the  briefly  sketched  cycle  of  development. 

Steenstrup's  work  on  the  cestodes  did  not  attract  the  same  atten- 
tion as  his  work  on  trematodes.  Steenstrup  always  insisted  on  the 
'^ nurse"  nature  of  the  cysticerci  and  other  bladder-worms.  Abildgaard 
(1790),  as  well  as  Creplin  (1829  and  1839),  had  already  furnished 
the    information    that   certain    sexless    cestodes    (Schistocephalns    and 


14  THE   ANIMAL   PARASITES   OF   MAN 

Ligtila)  from  the  abdomen  of  fishes  only  become  mature  after  their 
transference  to  the  intestine  of  aquatic  birds.  These  passive 
migrations  were  confirmed  in  an  entire  series  of  other  cestodes, 
particularly  by  v.  Siebold  (1844,  1848,  1850)  and  E.  J.  van  Beneden 
(1849),  not  by  actual  experiment^  but  by  undoubted  observation. 

It  was  correctly  believed  that  the  ova  or  oncospheres  penetrate 
into  certain  intermediate  hosts,  in  which  they  develop  into  unseg- 
mented  larvae.  Here  they  remain  until,  with  their  host,  they  are 
swallowed  by  some  predacious  animal.  They  then  reach  the  intestine, 
being  freed  from  the  surrounding  membranes  through  the  process 
of  digestion,  and  settle  themselves  there  to  form  the  adult  chain 
of  proglottides.  Though  some  few  scientists,  such  as  P.  J.  van 
Beneden  and  Em.  Blanchard,  deduced  from  these  observations  that 
the  bladder-worms  (Cysticerci),  which  had  hitherto  been  regarded  as 
a  separate  class  of  helminthes,  were  only  larval  Taeniae,  this  correct 
view  was  not  at  first  universally  accepted.  The  foundation  was  too 
slight,  and  van  Beneden  was  of  opinion  that  the  Cysticerci  were  not 
necessary,  but  only  appeared  incidentally. 

V.  Siebold  was  a  strenuous  opponent  to  this  theory,  notwith- 
standing his  experiences  on  the  change  of  hosts  of  the  Tetra- 
rhynchus.  Together  with  Dujardin  (1850)  he  conjectured  that  the 
Taeniae  underwent  a  deviating  cycle  of  development.  He  w^as  of 
opinion  that  the  six-hooked  oncospheres  left  the  lintestine,  in 
which  the  older  generation  lived,  and  were  scattered  about  with 
the  faeces,  and  finally  re-entered  per  os  (i.e.,  with  water  and  food) 
a  host  similar  to  the  one  they  had  left,  in  the  intestine  of  which 
they  w^ere  directly  transformed  into  tapew^orms.  A  change  of  host 
such  as  occurred  in  other  cestodes  was  not  supposed  to  take  place 
(the  history  of  the  cestodes  was  at  this  time  not  entirely  estab- 
lished). As  the  oncospheres  of  the  Taenia  are  enveloped  in  one 
calcareous  or  several  softer  coverings  which  they  cannot  leave 
actively,  and  as,  in  consequence  of  this  condition,  innumerable 
oncospheres  cannot  penetrate  into  an  animal,  and  others  cannot 
reach  the  proper  animal,  v.  Siebold  conceded,  at  least  for  the 
latter,  the  possibility  of  a  further  development.  But  this  was  only 
supposed  to  occur  because  they  had  either  invaded  wrong  hosts, 
or,  having  reached  the  right  hosts,  had  penetrated  organs  unsuit- 
able to  their  development,  and  had  thus  gone  astray  in  their 
travels,  and  had  become  hydropically  degenerated  taeniae.  This 
was  v.  Siebold's  explanation  of  bladder-worms.  Naturally,  v.  Siebold 
himself  conjectured  that  a  recovery  of  the  diseased  tapeworm  might 
occur,  in  a  few  exceptional  cases,  after  transmission  into  the  correct 
host,  as,  for  instance,  in  the  CysiicerciLS  fasciolaris  of  mice,  the  host  of 
which  is  the  domestic  cat,  and  in  which  there  is  a  seemingly  normally 


ORIGIN    OF    PARASITES 


15 


developed  piece  of  tapeworm  situated  between  the  caudal  vesicle  and 
the  cysticercus  head. 

Guided  by  correct  views,  F.  Kiichenmeister  undertook  in  Zittau 
the  task  of  confirming  the  metamorphosis  of  Cysticercus  pisiformis 
of  hares  and  rabbits,  into  tapeworms  in  the  intestine  of  the  dog 
by  means  of  feeding  experiments.  The  first  reports  on  the  subject, 
published  in  1851,  were  not  likely  to  meet  with  universal  approval, 
because  Kiichenmeister  first  diagnosed  the  actual  tapeworm  he  had 
been  rearing  as  Tcenia  crassiceps,  afterwards  as  Tcenia  serrata,  and 
finally  as  Tcenia  pisiformis  n.  sp.  However,  in  any  case,  Kiichenmeister, 
by  means  of  the  reintroduction  of  experimental  investigation,  rendered 
a  great  service  to  helminthology. 

The  publication  of  Kuchenmeister's  works  induced  v.  Siebold 
to  undertake  similar  experiments  (1852  and  1853),  which  were 
partly  published  by  his  pupil  Lewald  in  1852.  But  the  positive 
results  obtained  hardly  changed  Siebold's  opinion,  for  although  he 
no  longer  considered  the  bladder-worms  as  hydropically  degenerated 
tapeworms,  he  still  regarded  them  as  taeniae  that  had  strayed.  The 
change  of  opinion  was  partly  due  to  an  important  work  of  the 
Prague  zoologist,  v.  Stein  (1853).  He  was  able  to  examine  the 
development  of  a  small  bladder-worm  in  the  larvae  of  the  well-known 
meal-worm  {Tenebrio  molitor)  and  to  demonstrate  that,  as  Goeze  had 
already  proved  in  the  case  of  Cysticercus  fasciolaris  of  mice,  first  the 
caudal  vesicle  is  formed  and  then  the  scolex,  whereas  Siebold  believed 
that  in  bladder-worms  the  posterior  end  of  the  scolex  was  formed 
first,  and  that  this  posterior  end  underwent  a  secondary  hydropic 
degeneration. 

In  opposition  to  v.  Siebold,  Kiichenmeister  successfully  proved 
the  necessity  of  the  bladder- worm  stage  by  rearing  tapeworms 
in  dogs  from  the  Cysticercus  tenuicollis  of  domestic  mammals  and 
from  the  Ccenurus  cerebralis  of  sheep.  He,  and  simultaneously 
several  other  investigators  independently,  succeeded,  with  material 
provided  by  Kiichenmeister,  in  rearing  the  Ccenurus  cerebralis  in 
sheep  from  the  oncospheres  of  the  Tcenia  ccenurus  of  the  dog  (1854). 
R.  Leuckart  obtained  similar  results  in  mice  by  feeding  them  with  the 
mature  proglottides  of  the  Tcenia  crassicollis  of  cats  (1854). 

Kiichenmeister  also  repeatedly  reared  the  Taenia  solium  of  man 
from  the  Cysticercus  cellulosce  of  pigs  (1855),  and  from  the  embryos 
of  this  parasite  P.  ].  van  Beneden  succeeded  in  obtaining  the  same 
Cysticercus  in  the  pig  (1854).  As  Kiichenmeister  distinguished  the 
Tcenia  mediocanellalaj  known  to  Goeze  as  Tcenia  saginata,  amongst 
the  large  taeniae  of  man  (1851),  so  it  was  not  long  before  R.  Leuckart 
■(1862)  succeeded  in  rearmg  the  cysticercus  of  the  bookless  tapeworm 
in  the  ox.  It  is  particularly  to  this  last-named  investigator  that 
helminthology   is   indebted    more   than   to    any   other   author.      He 


l6  THE   ANIMAL   PARASITES   OF   MAN 

followed  the  gradual  metamorphosis  from  oncospheres  to  cystic 
worms  in  all  its  details. 

In  view  of  all  the  researches  that  were  made,  and  which  are  too 
numerous  to  mention  individually,  the  idea  that  bladder-worms  are 
abnormal  or  only  incidental  forms  had  to  be  abandoned.  Every- 
thing pointed  to  the  fact  that  in  all  cestodes  the  development  is 
divided  between  two  kinds  of  animals  ;  in  one — the  host,  the  adult 
tapeworm  is  found  ;  while  in  the  other,  the  intermediate  host,  we 
find  some  form  or  other  of  an  intermediate  stage  (cysticercus  in 
the  broadest  sense).  The  practical  application  of  this  knowledge  is 
self-evident.  If  no  infected  pork  or  beef  is  ingested,  no  tapeworm 
can  be  acquired,  and  also  the  rearing  of  cysticerci  in  the  human  body 
is  prevented  by  avoidmg  the  introduction  of  the  eggs  of  tapeworms. 

Though  these  results  were  definitely  proved  by  numerous 
researches,  yet  they  have  been  repeatedly  challenged,  notably  by 
J.  Knoch  (1862)  in  Petrograd,  who,  on  the  basis  of  experiments, 
sought  to  confirm  a  direct  development  without  an  intermediate  host 
and  ciliated  stage,  at  all  events  as  regards  Dibothriocephalus  laUis. 
However,  the  repeated  communications  of  this  author  met  with 
but  little  favour  from  competent  persons,  partly  because  the 
experiments  were  conducted  very  carelessly,  and  partly  because 
their  repetition  on  dog  and  man  (R.  Leuekart)  had  no  results 
(1863).  It  was  only  in  1883  that  Braun  was  able  to  prove  that  the 
developmental  cycle  of  Dibothriocephalus  latus  is  similar  to  that  of 
other  Cestodes.  The  results  obtained  in  other  places  by  Parona, 
Grassi,  Ijima  and  Zschokke  render  any  discussion  of  Kiichenmeister's 
conclusions  unnecessary.^  Long  after  Knoch,  a  French  author, 
P.  Megnin,  also  pleaded  for  the  direct  development  of  some 
cestodes,  and  especially  some  taeniae.  He  (1879)  also  sought  to 
prove  a  genetic  connection  between  the  bookless  and  armed  tape- 
worms of  mammals,  but  the  arguments  he  adduced,  so  far  as  they 
rest  on  observations,  can  be  easily  refuted  or  attributed  to  mis- 
interpretation. Only  one  of  these  arguments  is  correct,  namely, 
that  the  number  of  the  species  of  taeniae  with  which  we  are  acquainted 
is  far  larger  than  that  of  the  corresponding  cystic  forms ;  but  this 
disparity  alone  cannot  be  taken  as  a  proof  of  direct  development. 
It  can  only  be  said  that  our  knowledge  in  this  respect  is  deficient. 
As  a  matter  of  fact,  we  have  during  recent  years  become  acquainted 
with  a  large  number  of  cystic  forms,  hitherto  unknown,  belonging 
to  taeniae  which  have  long  been  familiar.  It  must  also  be  borne  in 
mind  that  no  man  in  his  lifetime  can  complete  an  examination  for 
bladder-worms  of  the  large  number  of  insects,  for  instance,   which 

'  Refer  to  the  collected  literature  under  Dibothriocephahis  latus,  and  the  reply  to  Klichen- 
meister  by  Braun  ('*  Ueber  den  Zwischenwirt  des  breit.  Bandw."     Wurzb.  :  Stuber,  1886). 


DEVELOPMENT   OP^   PARASITES  \*J 

may  destroy  an  entire  generation  of  an  insectivorous  species  of  bird 
within  a  small  district. 

Naturally  it  does  not  follow  that  direct  development  in  the 
cestodes  is  altogether  lacking.  The  researches  of  Grassi  (1889)  have 
furnished  an  example  in  Hyiiienolepis  {iceiiia)  imirina,  which  shows 
that  development  may  sometimes  take  place  without  an  intermediate 
host,  notwithstanding  the  retention  of  the  cystic  stage.  It  was  found 
that  the  oncospheres  of  this  species,  introduced  into  rats  of  a  certain 
age,  after  a  time  grow  into  tapeworms  without  leaving  the  intestine, 
but  not  directly,  for  they  bore  into  the  intestinal  wall,  where  they 
pass  the  cystic  stage,  the  cysts  afterwards  falling  into  the  intestinal 
lumen,  where  they  develop  into  tapeworms.  The  recent  experiments 
of  NicoU  (1911)  show  that  the  larval  stages  of  Hyinenolepis  uiurina 
also  occur  in  the  rat-flea,  Cera tophyllits  fasciatus. 

Important  observations  were  soon  made  on  the  remaining 
groups  of  helminthes.  The  discussion  on  the  origin  of  parasites 
soon  became  confined  to  the  helminthes.  Amongst  the  Nematoda, 
it  had  long  been  known  that  encapsuled  forms  existed  that  had 
at  first  been  regarded  as  independent  species,  but  very  soon  they  were 
pronounced  to  be  immature  forms,  in  consequence  of  their  lack 
of  sexual  organs.  Though  Dujardin  and  also  v.  Siebold  regarded 
them  as  ''strayed"  animals,  v.  Stein  (1853)  very  promptly  demon- 
strated that  the  progeny  of  the  nematodes  were  destined  to  travel 
by  discovering  a  perforating  organ  in  the  larval  nematodes  of  the 
mealworm.  This  was  first  experimentally  confirmed  (i860)  by 
R.  Leuckart,  R.  Virchow  and  Zenker,  all  of  whom  succeeded  not 
only  in  bringing  to  maturity  the  muscle  Trichinae  (known  since  1830) 
in  the  intestine  of  the  animals  experimented  upon,  but  were  likewise 
able  to  follow  the  migrations  of  the  progeny.  Of  course,  the 
encapsulating  brood  remained  in  the  same  organism,  and  in  this 
respect  deviated  from  the  broods  of  other  helminthes  which  escape 
into  the  outer  world  and  find  their  way  into  other  animals,  but 
the  encapsuled  nematodes  could  no  longer  be  regarded  as  the 
result  of  straying.  Subsequently,  R.  Leuckart  worked  out,  more 
or  less  completely,  the  history  of  the  development  of  numerous 
nematodes,  or  pointed  out  the  way  in  which  further  investigations 
should  be  made.  It  has  been  found  that  in  nematodes  far  more 
frequently  than  in  other  helminthes,  the  typical  course  of  develop- 
ment is  subject  partly  to  curtailment  and  partly  to  complications, 
which  sometimes  considerably  increase  the  difficulties  of  investigation 
and  have  hitherto  prevented  the  attainment  of  a  definite  conclusion, 
though  the  way  to  it  is  now  clear. 

In  a  similar  manner  the  works  of  R.  Leuckart  have  cleared 
up   the    development   of    the  Acanthocephala  and  Lingttatulida.     Of 


l8  THE   ANIMAL   PARASITES   OF   MAN 

course,  mucli  still  remains  to  be  done.  So  far,  we  do  not  even 
know  all  the  helminthes  of  man  and  of  the  domestic  animals 
in  all  their  phases  of  life,  and  still  less  is  known  of  those  of 
other  animals.  We  are  indebted  to  the  discoveries  of  the 
last  fifty  years  for  the  knowledge  arrived  at,  though  compara- 
tively few  names  are  connected  with  it.  The  gross  framework 
is  revealed,  but  the  gaps  have  only  been  filled  up  here  and 
there.  However,  we  may  trustfully  leave  the  completion  of  the 
whole  to  the  future,  without  fear  that  any  essential  alterations  will 
take  place. 

The  deductions  to  be  drawn  are  as  follows  :  That  the  helminthes 
like  the  ectoparasites  multiply  by  sexual  processes,  that  the  entire  course 
of  development  of  the  helminthes  is  rarely  or  never  gone  through 
in  the  same  host  as  is  the  case  with  several  ectoparasites,  that  the 
progeny  at  an  earlier  or  later  stage  of  development,  as  eggs,  embryos, 
or  larvae,  quit  the  host  inhabited  by  the  older  generation,  and  almost 
always  attain  the  outer  world  :  only  in  Trichinella  does  the  develop- 
ment take  place  directly  in  the  definite  host.  Where  the  eggs  have 
not  yet  developed  they  go  through  the  embryonic  evolution  in  the 
outer  world.  The  young  larvae  are  transmitted,  either  still  enclosed 
within  the  Qgg  or  embryonic  covering,  to  the  intermediate  host  or 
more  rarely  they  are  transferred  straight  to  the  final  host.  In  other 
cases  they  may  hatch  out  from  their  envelopes,  and  after  a  longer  or 
shorter  period  of  free  life,  during  which  they  may  partake  of  food  and 
grow%  they,  as  before,  penetrate,  usually  in  an  active  way,  into  an 
intermediate  host,  or  at  once  invade  the  final  host.  Exceptionally 
{e.g.f  Rhabdonema),  during  the  free  life  there  may  be  a  propagation  of 
the  parasitic  generation,  and  in  this  case  only  the  succeeding  genera- 
tion again  becomes  parasitic,  and  then  at  once  reaches  its  final  host. 
The  young  forms  which  have  invaded  the  final  host  become  mature  in 
the  latter,  or  after  a  longer  or  shorter  period  of  parasitism  again  wander 
forth  (as  the  CEstridae,  Ichneumonidae,  etc.),  and  reach  the  adult  stage 
in  the  outer  world.  The  young  stages,  during  which  the  parasites 
undergo  metamorphoses  or  are  even  capable  of  producing  one  or 
several  intermediate  generations,  are  passed  in  the  intermediate  hosts 
until,  as  a  rule,  they  are  passively  carried  into  the  final  host  and  there 
complete  their  cycle  of  development  by  the  formation  of  the  organs 
of  generation.  This  mode  of  development,  the  spending  of  life  in 
two  different  kinds  of  animals  (intermediate  and  final  host),  is  typical 
of  the  helminthes.  This  is  manifested  in  the  Acanthocephala,  the 
Cestoda,  the  majority  of  the  endoparasitic  Trematoda,  a  number  of 
the  Nematoda,  and  the  Linguatulidae.  There  are  now  and  then 
exceptions,  however,  in  which,  for  instance,  the  host  and  intermediate 
host  change  order  {Trichinella y  Hymenolepis  miirina). 


DERIVATION   OF   PARASITES  I9 

Parasites  are  hardly  ever  inherited  amongst  animals.^  According 
to  a  few  statements,  however,  Trlcliinella  and  Cceniirus  are  supposed 
to  be  transmissible  from  the  infected  mother  to  the  foetus.  Other- 
wise most  animals  acquire  their  parasites,  especially  the  Entozoa,  from 
without,  the  parasites  penetrating  either  actively,  as  in  animals  living 
in  the  water,  or  passively  with  food  and  drink.  A  particular  pre- 
disposition to  worms  is  not  more  likely  than  a  spontaneous  origin 
of  parasites. 

Derivation  of  Parasites. — Doubt  now  no  longer  exists  as  to  the 
derivation  of  the  temporary  and  of  many  of  the  stationary  ecto- 
parasites from  free-living  forms.  This  conclusion  is  founded  on  the 
circumstance  that  not  only  are  there  numerous  intermediate  degrees 
in  the  manner  of  living  and  feeding  between  predacious  and  parasitic 
animals,  but  that  there  is  more  or  less  uniformity  in  their  structure. 
The  differences  that  exist  are  easily  explained  as  consequences  of 
altered  conditions  of  life.  The  case  is  more  difficult  in  regard  to 
groups  that  are  exclusively  parasitic  (Cestoda,  Treniatoda,  Acantho- 
cephala,  Liiiguatiilidce,  and  Sporozoa),  or  groups  that  are  chiefly 
parasitic  {Nematoda),  because  in  these  cases  the  gulf  that  divides 
these  forms  from  free-living  animals  is  wider.  It  is  true  that  we  know 
that  the  nearest  relatives  of  the  Lingtiatuiidoi  are  found  amongst  the 
Arachnoidea,  and  indeed  in  the  Acarlna ;  that,  moreover,  the  structure 
and  development  of  the  Sporozoa  refers  them  to  the  Protozoa^  and 
allows  some  of  them  to  be  regarded  as  the  descendants  of  the  lowest 
Rhlzopoda,  We  know  that  the  Trematoda,  and  through  these  the 
Cestoda,  are  closely  related  to  the  Turbellaria,  from  which  they  may 
be  traced.  The  Neinatoda,  and  still  more  the  Acanthocephala,  stand 
apart.  This  is  less  evident,  how^ever,  in  the  Nematoda,  for  there 
are  numerous  free-living  members  of  these  from  which  it  is  possible 
that  the  parasitic  species  may  be  descended.  Indeed,  this  seems 
more  than  probable  if  such  examples  as  Leptodera,  Rliabdonema  and 
Strongyloides  are  taken  into  consideration,  as  well  as  the  conditions 
of  life  of  free-living  nematodes.  These  mostly,  if  not  exclusively, 
spend  their  lives  in  places  where  decomposing  organic  substances 
are  present  in  quantities  ;  some  species  attain  maturity  only  in  such 
localities,  and  there  propagate  very  rapidly.  Should  the  favourable 
conditions  for  feeding  be  changed,  the  animals  seek  out  other 
localities,  or  they  remain  in  the  larval  form  for  some  time  until  more 
favourable  conditions  set  in.  It  is  comprehensible  that  such  forms 
are  very  likely  to  adopt  a  parasitic  manner  of  life  which  at  first  is 
facultative  {Leptodera,  Angnillula),  but  may  be  regarded  as  the  tran- 

'  However,  in  the  Protozoa  there  are  examples  of  hereditary  transmission  of  parasites,  e.g., 
in  the  case  of  Babesia  {Piroplasma)  bovis  and  Babesia  cants  in  their  invertebrate  hosts  (ticks) ; 
in  Crithidia  melophagia  and  Crithzdia  hyalommce  ;  and  in  the  case  of  SpirocJueta  diittoni  in  its 
invertebrate  host  (a  tick). 


20  THE   ANIMAL   PARASITES   OF   MAN 

sition  to  true  parasitism.  The  great  advantages  attached  to  a  parasitic 
Hfe  consist  not  only  in  protection,  but  also  in  the  supply  of  suitable 
food,  and  consequently  in  the  easier  and  greater  production  of  eggs, 
and  thus  fully  account  for  the  gradual  passage  of  facultative  parasitism 
into  true  parasitism.  In  many  forms  the  young  stages  live  free  for 
some  time  {Strongylidce),  in  others,  as  is  the  case  in  Rhabdoiieina, 
parasitic  and  free-living  generations  alternate  ;  in  others,  again,  the 
free  period  is  limited  to  the  agg  stage  or  entirely  suppressed. 

Though  it  is  possible  thus  to  connect  the  parasitic  with  the  free- 
living  nematodes,  by  taking  their  manner  of  life  into  account,  this 
matter  presents  greater  difticulties  in  regard  to  other  helminthes.  It 
is  true  that  the  segmented  Cestoda  may  be  connected  with  and  traced 
from  the  less  known  and  interesting  single-jointed  Cestoda  {Ainphilina, 
Archigetes,  Caryophyllc^us,  Gyrocotyle).  Trematodes  are  all  parasites, 
with  the  exception  of  one  group,  Temnocephaiidiv,  several  genera  and 
species  of  which  live  on  the  surface  of  the  bodies  of  Crustacea  and 
turtles  of  tropical  and  sub-tropical  freshwaters.  Teriinocephalid^^  nre, 
nevertheless,  predacious.  They  feed  on  Infusoria,  the  larvae  of  small 
insects  and  Crustacea.  So  far  as  is  known  they  do  not  nourish  them- 
selves on  part  of  the  host.  They  belong  to  the  group  of  commensals, 
or  more  correctly,  to  that  of  the  space  parasites,  which  simply  dwell 
with  their  host  and  do  not  even  take  a  portion  of  the  superfluity  of 
its  food.  However,  space  parasitism  may  still  be  regarded  as  the 
first  stage  of  commensalism,  which  is  again  to  be  regarded  as  a  sort 
of  transition  to  true  parasitism. 

It  is  possible  that  parasitism  came  about  in  this  way  in  the 
trematodes,  in  which  connection  we  must  first  consider  the  turbellaria- 
like  ancestors  of  the  trematodes.  Much  can  be  said  in  favour  of 
such  a  genetic  relationship  between  turbellaria  and  trematodes,  and 
.hardly  anything  against  it.  It  should  also  be  remembered  that 
amongst  the  few  parasitic  turbellaria  there  are  some  that  possess 
clinging  discs  or  suctorial  pores,  and  these  are  only  differentiated 
from  ectoparasitic  trematodes  by  the  possession  of  a  ciliated  integu- 
ment, which  is  found  only  in  the  larval  stages  of  the  latter. 

The  Acanthocephala  occupy  an  isolated  position.  Most  authors 
certainly  regard  them  as  related  to  the  nematodes ;  in  any  case, 
the  connection  is  not  a  close  one,  and  the  far-reaching  alterations 
which  must  have  occurred  prevent  a  clear  view.  Perhaps  the  free 
original  forms  of  Acanthocephala  are  no  longer  in  existence,  but 
that  such  must  have  existed  is  a  foregone  conclusion. 

An  explanation  of  the  change  of  host  so  frequent  in  parasites 
is  more  difficult  than  that  of  their  descent.  R.  Leuckart  is  of  opinion 
that  the  present  intermediate  hosts,  which  belong  principally  to  the 
lower  animals,  were  the  original  hosts  of  the  parasites,  and  fostered 


DERIVATION    OF   PARASITES  21 

both  their  larval  and  adult  stages.  It  was  only  in  course  of  time  that 
the  original  hosts  sank  to  the  position  of  intermediate  hosts,  the  cause 
for  this  alteration  being  that  the  development  of  parasites,  especially 
of  the  helminthes,  through  further  development  and  differentiation 
extended  over  a  larger  number  of  stages.  The  earlier  stages  remained 
in  their  original  hosts,  but  the  later  stages  sought  out  other  hosts 
(higher  animals).  To  prove  this,  Leuckart  points  out  that  the  mature 
stages  of  the  helminthes,  with  but  few  exceptions,  occur  only  in  the 
vertebrates  which  appeared  later  in  the  development  of  the  animal 
kingdom,  while  the  great  majority  of  intestinal  worms  of  the  lower 
animals  only  represent  young  stages,  which  require  transmission  into 
a  vertebrate  animal  before  they  can  become  mature.  The  few 
helminthes  that  attain  maturity  in  the  lower  animals  (Aspidogaster, 
Archigetes)  are  therefore  regarded  by  Leuckart  as  primitive  forms, 
and  he  compares  them  with  the  developmental  stages  of  helminthes, 
Aspidogaster  with  rediae,  Archigetes  with  cystic ^rcoids.  He  classes 
the  nematodes  that  become  mature  in  the  invertebrates  with  Angiiil- 
Inlidce,  i.e.,  with  saprophagous  nematodes  from  which  the  parasitic 
species  descend. 

Leuckart  therefore  regards  the  change  of  hosts  as  secondary, 
so  does  Sabatier.  The  latter,  however,  adduces  other  reasons  for 
this  (lack  of  clinging  organs  and  the  necessity  to  develop  them  in  an 
intermediary  stage)  ;  but  in  this  connection  he  only  considers  the 
Cestoda.  In  opposition  to  Leuckart,  R.  Moniez,  however,  is  con- 
vinced that  the  migrations  of  the  helminthes,  as  well  as  the  system 
of  intermediate  hosts,  represent  the  original  order  of  things.  Moniez 
tiaces  all  Entozoa  from  saprophytes,  but  only  a  few  of  these  were 
able  to  settle  directly  in  the  intestine  and  there  continue  their 
development.  These  are  forms  that  at  the  present  day  still  lack 
an  intermediate  host,  such  as  Trichocephalus,  Ascaris,  and  Oxyurls. 
In  most  other  cases  the  embryos,  however,  consisted  of  such  sapro- 
phytes as  were,  in  other  respects,  suitable  to  become  parasites,  but 
were  incapable  of  resisting  the  mechanical  and  chemical  influences 
of  the  intestinal  contents.  They  were  therefore  obliged  to  leave  the 
mtestine  at  once,  and  accomplished  this  by  penetrating  the  intestinal 
walls  and  burrowing  in  the  tissues  of  their  carriers.  In  this  position, 
assisted  by  the  favourable  conditions  of  nutrition,  they  could  attain 
a  relatively  high  degree  of  development.  Mechanical  reasons  pre- 
vented a  return  to  the  intestines,  where  the  eggs  could  be  deposited. 
Most  of  them  doubtless  died  off  as  parasites,  as  also  their  young 
stages  do  at  present  when  they  penetrate  WTong  hosts.  Some  of 
them,  nevertheless,  passively  reached  the  intestine  of  beasts  of  prey. 
Many  were  destroyed  in  the  process  of  mastication ;  for  a  small  part, 
however,  there  was  the  chance  of  reaching  the  intestine  of  a  beast 


22  THE   ANIMAL   PARASITES   OF   MAN 

of  prey  undamaged,  and  there,  having  become  larger  and  more 
capable  of  resistance,  maturity  was  attained.  By  means  of  this  in- 
cidental coincidence  of  various  favourable  circumstances,  these  pro- 
cesses, according  to  Moniez,  have  been  established  by  heredity  and 
Iiave  become  normal. 

This  is  not  the  place  to  express  an  opinion  either  for  or  against 
the  various  hypotheses  advanced,  but  the  existence  of  these  diametri- 
cally opposed  views  alone  will  show  the  great  difficulty  of  the 
question.  Independently,  however,  it  appears  more  natural  to  come 
to  the  conclusion  that  parasitism,  as  well  as  change  of  hosts,  were 
gradual  transitions. 

As  a  conclusion  to  this  introductory  chapter,  a  list  of  some  of 
the  most  important  works  on  the  parasitology  of  man  and  animals 
is  appended. 

LITERATURE. 

GOEZE,  J.  A.  E.  Versuch  einer  Naturgeschichte  der  Eingeweidewiirmer  thieri- 
scher  Korper.     Blankenburg,   1782.     4to,  471  pp.,  with  44  plates. 

Zeder,  J.  G.  H.  Erster  Nachtrag  zur  Naturgeschichte  der  Eingeweidewiirmer. 
von  J.  A.  E.  Goeze.     Leipzig,  1800.     4to,  with  6  tables. 

RUDOLPHI,  C.  A.  Entozoorum  sive  vermium  intestinalium  historia  naturalis. 
I,  Amstelod.,  1808;  ii,  i8og.     8vo,  with  18  plates. 

RUDOLPHi,  C.  A.     Entozoorum  synopsis.     Berol.,  1819.     8vo,  with  3  plates. 

Bremser,  J.  G.  Ueber  lebende  Wiirmer  im  lebenden  Menschen.  Wien,  1819. 
8vo,  with  4  plates. 

Bremser,  J.  G.  Icones  helminthum,  systema  Rudolphii  entozoologicum  illus- 
trantes.     Viennae,   1824.     Fol.   (Paris,    1837). 

DUJARDIN,  F.  Histoire  naturelle  des  helminthes  ou  vers  intestinaux.  Paris, 
1845.     8vo,  with  12  plates. 

DiESING,  C.  M.  Systema  helminthum.  2  vols.  Vindobonnae,  1850,  185 1.  8vo. 
Supplements  by  the  same  author  :  Revision  der  Myzhelminthen  (Report 
of  the  Session  of  the  Imp.  Acad,  of  Science.  Wien,  xxxii,  1858) ;  with 
addendum  (ibid.,  xxxv,  1859);  Revision  der  Cephalocotyleen  (ibid.,  xlix, 
1864,  and  xlviii,  1864) ;  Revision  der  Nematoden  (ibid.,  xlii,  1861) ;  Supple- 
ments (ibid.,  xliii,   1862). 

Beneden,  p.  J.  VAN.  Memoire  sur  les  Vers  intestinaux.  Paris,  1858.  4to,  with 
12  plates. 

KtJCHENMElSTER,  F.  Die  in  und  an  dem  Korper  des  lebenden  Menschen  vorkom- 
menden  Parasiten.     Leipzig,  1855.     8vo,  with  14  plates. 

Leuckart,  R.  Die  menschlichen  Parasiten  und  die  von  ihnen  herriihrenden 
Krankheiten.     I,  Leipzig,  1863;  II,  Leipzig,  1876.     8vo. 

COBBOLD,  T.  Sp.  Entozoa ;  an  Introduction  to  the  Study  of  Helminthology. 
London,  1864.     8vo.     Supplement,  London,  1869. 

Davaine,  C.  Traite  des  entozoaires  et  des  maladies  vermineuses  de  I'homme  et 
des  animaux  domestiques.     2nd  edit.     Paris,    1877.     8vo. 

LiNSTOW,  O.  V.  Compendium  der  Helminthologie,  ein  Verzeichniss  der 
bekannten  Helminthen,  die  frei  oder  in  thierischen  Korpern  leben, 
geordnet  nach  ihren  Wohnthieren,  unter  Angabe  der  Organe,  in  dencn 
sie  gefunden  sind,  und  mit  Beifiigung  der  Litteraturquellen.  Hanov., 
1878.     8vo.     Supplement,  including  the  years  1878-1888,  Hanov.,   1888. 

COBBOLD,  T.  Sp.  Parasites;  a  Treatise  on  the  Entozoa  of  Man  and  Animals, 
including  some  Account  of  the  Entozoa.     London,  1879.     8vo. 

Leuckart,  R.  Die  Parasiten  des  Menschen  und  die  von  ihnen  herriihrenden 
Krankheiten.  2nd  edit.  Leipzig,  1879-1886.  The  Protozoa,  Cestodes, 
Trematodes  and  Hirudinea  have  hitherto  appeared  (continued  by 
Brandes). 

BtJTSCHLi,  O.  Protozoa  in  Bronn's  Klass.  u.  Ordn.  d.  Thierreichs.  Vol.  i, 
Leipzig,  1 880-1889.     8vo,  with  79  plates. 


LITERATURE 


23 


Braun,  M.  Trematodes  in  Bronn's  Klass.  u.  Ordn.  d.  Thierreichs.  Vol.  iv,  i, 
Leipzig,  1879-1893.  8vo,  with  33  tables.  (The  first  thirteen  sheets,  com- 
prising the  history  of  the  worms  up  to  1830,  were  compiled  by  H. 
Pagenstecher.) 

ZlJRN,  F.  A.  Die  thierischen  Parasiten  auf  und  in  dem  Korper  unserer  Haus- 
saugethiere,  sowie  die  durch  erstere  veranlassten  Krankheiten,  deren 
Behandlung  und  Verhiitung.  2nd  edit.  Weimar,  1882.  8vo,  with  4 
plates. 

COBBOLD,  T.  Sp.  Human  Parasites;  a  Manual  of  Reference  to  all  the  Known 
Species  of  Entozoa  and  Ectozoa.     London,  1882.     8vo. 

KUCHENMEISTER,  F.,  and  F.  A.  ZURN.  Die  Parasiten  des  Menschen.  2nd  edit. 
Leipzig,  1888.,    8vo,  with  15  plates. 

Blanchard,  R.     Traite  de  zoologie  medicale.     I,  Paris,  1889;  II,  i8go,     Svo. 

Neumann,  L.  G.  Traite  des  maladies  parasitaires  non  microbiennes  des 
animaux  domestiques.  2nd  edit.  Paris,  1892.  Svo.  English  edit.,  trans- 
lated by  G.  Fleming.  •  2nd  edit.,  revised  by  J.  Macqueen.  1905.  London  : 
Bailliere,  Tindall  and  Cox. 

Looss,  A.     Schmarotzerthum  in  der  Thierwelt.     Leipzig,  1892.     Svo. 

Railliet,  a.  Traite  de  zoologie  medicale  et  agricole.  2nd  edit.  I,  Paris, 
1S95.     Svo. 

ParoN!\,  C.  L'elmintologia  italiana  da'  suoi  primi  tempi  all'  anno  1890. 
Genova,    1894.     Svo. 

Braun,  M.  Cestoda  in  Bronn's  Klass.  u.  Ordn.  d.  Thierreichs.  Vol.  iv,  2, 
Leipzig,  1S94-1900.     Svo,  with  24  plates. 

MOSLER,  F.,  and  E.  Peiper.  Thier  Parasit.  (Spec.  Path.  u.  Ther.  v.  H. 
Nothnagel.     Vol.   vi.)     Wien,  1894.     Svo,  with   124  illustrations. 

Laveran^  a.,  et  R.  Blanchard.  Les  hematozoaires  de  I'homme  et  des  anim, 
Paris,    1895.     i2mo,    with   30  figs. 

Sluiter,  C.  R.     De  dierl,   paras,   v.   d.   mensch  en  van  onze  huisdier.     Haag, 

1895.  Svo. 

Blanchard,  R.     Malad.  parasit.,  paras,   animaux,  paras,  veget.  a  I'exclus.  des 

bacter.      (Traite  de  pathol.  gen.   de  Ch.  Bouchard,  vol.  ii.)     Paris,  1S95. 

Svo,  with  70  figs. 
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With    Supplement,     1898,    and    continued    as    Bibl.     d.    klin.     Entomol. 

Miinchen,   1S99-1900. 
MONIEZ,  R.     Traite  de  parasitol.  anim.   et  veget.  appl.  a  la  medecine.     Paris, 

1896.  Svo,  with  116  figs. 

Weichselbaum.     Parasitologie    (Weil's    Handb.    d.    Hyg.).     Jena,    1S9S.      Svo, 

with  78  illustrations, 
Kraemer,  a.     Die  thierischen   Schmarotzer  des  Auges   (Grafe  and  Samische's 

Handb.  d.  ges  Augenheilk.).     Leipzig,  1S99.     Svo,  with  16  illustrations. 
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(atlas  with  15  plates). 
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Zoology.     AVashington,   1902  (U.S.   Dept.  of  Agric,  Bur.  of  Anim.  Ind., 

Bull.  No.  39). 
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animaux.     4  e  edit.     Paris,  1911. 
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222  Abb, 
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parasitologique.     Paris,  1906.     With  500  figs. 
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gart, 1904. 
Stiles,  Ch.  W.     The  International  Code  of  Zoological  Nomenclature  as  applied 

to  Medicine  (Hygienic  Lab.,  Bull.   No.  24,  Washington,   1905)- 
Stiles^  C.  W.,  and  Hassall,  A.     Trematoda  and  Trematode  Diseases.     (Index 

Catalogue  of   Med.    and  Vet.    Zoology.)     Hygienic  Lab.,    Bull.    No.    ZT^ 

Washington,   1908. 


24  THE   ANIMAL    PARASITES   OF   MAN 

Stiles,   C.    \\.,    and   Hassall,    A.     Cestoda   and    Cestodaria.      Hygienic    Lab., 

BulL  No.  85,  Washington,  1Q12. 
Laloy,    L.     Parasitisme    et    mutualisme    dans    la    nature.     Paris,     igo6.     8vo, 

284  pp.,  82  figs. 
Theobald,   F.  V.     A  Monograph  of  the  Culicidae  of  the  World.     5  vols,   and 

plates.      iQoi-igio.     London  :  Brit.   Museum,  Nat.  Hist. 
James^    S.    p.,    and    LlSTON^    W.    G.     The    Anopheline    Mosquitoes    of    India. 

2nd  edit.      igii.     Calcutta  :  Thacker,  Spink  and  Co. 
Howard,  L.    O.,   Dyar^  H.    G.,   and  Knab,   F.     The   Mosquitoes  of  North   and 

Central    America   and    the    West    Indies.     2    vols.      1Q12.     Washington  : 

Carnegie   Institution. 
Austen,  E.  E.     African  Blood-sucking  Flies,      iqoq.     London  :    Brit.   Museum, 

Nat.  History. 
Austen^  E.   E.     A   Handbook  of  Tsetse-flies,      igii.     London:    Brit.   Museum, 

Nat.   History. 
Castellani,  a.,  and  Chalmers^  A.  J.     Manual  of  Tropical  Medicine.    2nd  edit. 

1,747  pp.      igi3.     London  :   Bailliere,  Tindall  and  Cox. 
KOLLE  and  Wassermann.     Handbuch  der  pathogenen  mikroorganismen.    Jena  : 

Gustav  Fischer. 
MiNCHIN,     E.     A.     An    Introduction    to    the     Study    of    the    Protozoa.       igi2. 

London  :  Arnold. 
Laveran,    a.,    et    Mesnil,    F.     Trypanosomes   et    Trypanosomiases.     2nd    edit. 

igi2.     Paris  :   Masson  and  Co. 
DOFLEIN^   F.     Lchrbuch  der   Protozoenkunde.     3rd  edit.      igii.     Jena:    Gustav 

Fischer. 
NuTTALL^  G.   H.  F.,  Warburton,  C,  Cooper,  W.  F.,  and  Robinson^  L.   E. 

Ticks — a   Monograph   of    the    Ixodoidea.     Pt.    I    (igo8).     Pt.    II.    (igii). 

University  Press,  Cambridge,  England. 
Brumpt,  E.     Precis  de  Parasitologie.     2nd  edit.      igi3.     Paris  :  Masson  and  Co. 
Patton,  W.  S.,  and  Cragg,  F.  W.    A  Text-book  of  Medical  Entomology.     igi3. 
Christian  Literature  Society  of  India  :  London,  Madras,  and  Calcutta. 

JOURNALS. 

For  current  researches  the  following,  among  others,  should  be  consulted  :  — 

Annals  of  Troi)ical  Medicine  and  Parasitology^  Liverpool. 

Annales  de  VInstitut  Pasteur,  Paris, 

Archives  de  Parasitologie,  Paris. 

Archives  de  Zoologie  Exferimentale  et  Generate,  Paris. 

ArchiiJ   fiir  Protistenhunde,  Jena. 

Archiv  fiir  Schiffs-  itnd  Tro-pen-Hygiene,  Leipzig. 

Bulletin  of  Entomological  Research,  London. 

Bulletin  de  VInstitut  Pasteur^  Paris. 

Bulletin  de  la  Societe  de  Pathologic  Exotique,  Paris. 

Bulletins  of  the  Bureau  of  Animal  Industry^  Washington. 

Centralhlatt  fiir  Bakteriologie  und  Parasitenkunde,  Jena. 

Com  ft.  Rend.  Acad.  Sci.,  Paris. 

Comft.  Rend.  Soc.  Biol.,  Paris. 

Indian  Journal  of  Medical  Research^  Calcutta. 

Journal  of  Ex-perimental  Medicine,  New  York. 

Journal  of  Medical  Research^  Boston. 

Memorias  do  Instituto  Osivaldo  Cruz,  Rio  de  Janeiro. 

Parasitology,  Cambridge. 

Proceedings  of  the  Royal  Society,  London. 

Quarterly  J ournal  of  Microscopical  Science,  London. 

Review  of  Applied  Entomology,  London. 

Tropical  Diseases  Bulletin  (London  :  Tropical  Diseases  Bureau). 

Zeitschrift  fiir  hifehtionshranhheiten,   Berlin. 


THE  ANIMAL  PARASITES  OF  MAN 


Man  is  one  of  those  organisms  in  or  on  which  a  whole  host  of  parasites  find 
conditions  suitable  for  their  existence:  Protozoa,  Platyhelminihes,  Nematoda, 
Acanthocephala,  Hirudinea,  and  a  large  number  of  Arthropoda  (Arachnida  as  well 
as  Insects)  all  include  members  which  are  parasites  of  man.  These  animals  either 
live  on  the  external  surface  of  the  body  or  within  the  intestine  and  its  appendages. 
Other  organs  and  systems  are  not  quite  free  from  foreign  organisms — we  are 
acquainted  with  parasites  in  the  skeletal  system,  in  the  circulatory  system,  in  the 
brain,  in  the  muscles,  in  the  excretory  and  genital  organs,  and  even  in  the  organs  of 
sense. 

It  is  possible,  and  perhaps  might  be  advantageous,  to  arrange  and  describe  the 
parasites  of  man  according  to  the  situations  in  which  they  are  found  (parasites  of  the 
skin,  intestinal  parasites,  etc.).  Their  description  in  the  various  stages  of  develop- 
ment would,  however,  be  disturbed  when,  as  is  generally  the  case,  the  different  stages 
are  passed  in  different  organs,  and  a  work  which  treats  more  fully  of  the  natural 
history  of  the  parasites  than  of  the  local  disorders  to  which  they  give  rise  would 
suffer  thereby.  It  is,  therefore,  preferable  to  describe  the  parasites  of  man  in  their 
systematic  order,  and  to  mention  their  different  situations  in  man  in  describing  each 
species. 

A.     PROTOZOA,     . 

BY 

H.    B.    FANTHAM,  M.A.,  D.Sc. 

All  those  animal  organisms  which  throughout  their  entire  life  never  rise  above 
the  unicellular  stage,  or  merely  form  simple,  loose  colonies  of  similar  unicellular 
animals,  are  grouped  under  the  term  Protozoa  (Goldfuss,  1820),  as  the  simplest  types 
of  animal  life.  All  the  vital  functions  of  these,  the  lowest  forms  of  animals,  are 
carried  out  by  their  body  substance,  the  protoplasm  (sarcode).  Often  particular 
parts  possess  special  functions,  but  the  limits  of  a  cell  are  never  over-stepped 
thereby.  These  special  parts  of  the  cell  are  called  "  cell-organs  "  ;  recently  they 
have  been  termed  "  organella?." 

The  living  protoplasm  has  the  appearance  of  a  finely  granular,  viscid  substance 
which,  as  a  -rule,  when  not  surrounded  by  dense  investing  membranes  or  skeletons, 
exhibits  a  distinct  kind  of  movement,  which  has  been  termed  amoeboid.  According 
to  the  species,  processes  of  different  forms  and  varying  numbers  called  pseudopodia 
are  protruded  and  withdrawn,  and  with  their  assistance  these  tiny  organisms  glide 
along — it  might  almost  be  said  f^ow  along— over  the  surface.  In  most  Protozoa  two 
layers  of  cytoplasm  may  be  recognised,  and  distinguished  by  their  appearance  and 
structure,  namely,  the  superficially  situated,  viscid,  and  quite  hyaline  ectosarc  or 
ectoplasm,  and  the  more  fluid  and  always  granular  endosarc  or  endoplasm,  which 
is  entirely  enveloped  by  the  ectoplasm.  The  two  layers  have  different  functions  ;  the 
movements  originate  from  the  ectoplasm,  which  also  undoubtedly  fulfils  the  functions 
of  breathing,  introduction  of  food  and  excretion.  The  endoplasm,  which  in  some  forms 
(Radiolaria)  is  separated  from  the  ectoplasm  by  a  membrane,  undertakes  the  digestion 


26  THE   ANIMAL   PARASITES   OF   MAN 

of  the  food.  To  this  distribution  of  functions  between  the  various  layers  ot  cyto- 
plasm is  due  the  development  of  particular  cellular  organs,  such  as  the  appearance 
of  cilia,  flagella,  suctorial  tubules  (in  the  Suctoria)  and  the  myophan  striations,  which 
are  contractile  parts  of  the  ectoplasm  in  Infusoria  and  Gregarines.  In  many  cases 
(Flagellata,  Ciliata),  an  area  is  differentiated  for  the  ingestion  of  food  (oral  part, 
cytostome)  to  which  there  is  often  added  a  straight  or  curved  opening  (cytopharynx), 
through  which  the  food  reaches  the  endoplasm.  The  indigestible  residue  is  either  cast 
off  through  the  oral  part  or  excreted  by  a  special  anal  part  (cytopyge).  In  rare 
cases,  structures  sensitive  to  light,  the  so-called  pigment  or  eye  spots  are  developed, 
e.g.^  Eugle7ta.  In  the  case  of  Infusoria  the  endoplasm  circulates  slowly,  and 
agglomerations  of  fluids  (food  vacuoles)  sometimes  appear  around  each  bolus  of 
food  ;  in  these  vacuoles  the  food  is  digested  under  the  action  of  certain  materials 
(ferments).  Even  in  the  lowliest  Protozoa  fluids  to  be  excreted  are,  as  a  rule, 
gathered  into  one,  or,  more  rarely,  several  contractile  vacuoles,  which  regularly 
discharge  their  contents.  This  action,  however,  is  to  a  certain  extent  governed 
by  the  temperature  of  the  surrounding  medium.  In  some  Infusoria  a  tube-like 
channel  in  the  cytoplasm  is  joined  to  the  contractile  vacuole  which  usually  occupies 
a  certain  position  ;  this  forms  a  sort  of  excretory  duct,  and  there  are  also  supply- 
canals  leading  to  these  organellae. 

Very  frequently  various  substances  are  deposited  in  the  endoplasm,  such  as  fatty 
granules,  drops  of  oil,  pigment  granules,  bubbles  of  gas  or  crystals.  More  solid 
skeletal  substances  are  secreted  in  or  on  the  ectoplasm.  To  the  latter  belong  the 
cuticle  of  the  Sporozoa  and  Infusoria,  the  chalky  shells  containing  one  or  several 
chambers  of  the  Foraminifera,  the  siliceous  and  very  ornamental  framework  of  the 
Radiolaria,  and  the  chitinous  coat  of  many  Flagellata,  Infusoria,  etc.  Some  forms 
make  use  of  foreign  bodies  found  in  their  surroundings,  such  as  grains  of  sand,  to 
construct  their  protective  coverings. 

The  food  often  consists  of  small  animal  or  vegetable  organisms  and  of  organic 
waste  ;  it  is  usually  introduced  in  toto  into  the  endoplasm.  On  the  other  hand,  the 
Suctoria  extract  nourishment  from  their  prey  by  means  of  their  tentacles.  Many 
parasitic  species  also  ingest  solid  food,  others  feed  by  endosmosis. 

In  all  cases  one  nucleus  at  least  is  present.  It  is  true  that  the  existence  of  non- 
nucleated  Protozoa,  the  so-called  Mofiera,  is  still  insisted  upon,  but  some  of  these 
have  already  proved  to  be  nucleated,  and  the  presence  of  nuclei  in  the  others  will  no- 
doubt  be  established.  Very  often  the  number  of  nuclei  increases  considerably,  but 
these  multinucleate  stages  are  always  preceded  by  uninucleate  stages.  In  the 
Infusoria,  in  addition  to  the  larger  or  principal  nucleus  (macronucleus)  there  is 
usually  a  smaller  reproductive  nucleus  (micronucleus).  This  dualism  of  the  nuclear 
apparatus  is  considered  by  some  to  be  general,  and  usually  to  appear  first  at  the 
onset  of  reproduction. 

The  form  and  structure  of  the  nucleus  vary  greatly  in  different  species.  There 
are  elongate,  kidney-shaped,  or  even  branched  nuclei  as  well  as  spherical  or  oval 
ones.  In  addition  to  vesicular  nuclei  with  a  distinct  karyosome  and  incidentally 
also  with  a  nuclear  membrane,  homogeneous  and  more  solid  formations  are 
frequently  encountered.  The  nuclei  are  always  differentiated  from  the  protoplasm 
by  their  reactions,  particularly  in  regard  to  certain  stains. 

In  many  Protozoa  an  extra-nuclear  mass,  sometimes  compact,  sometimes  diffuse,, 
arises  from  or  near  the  nucleus.  This  mass,  whose  staining  reactions  resemble  those 
of  the  nucleus,  is  termed  the  chromidial  apparatus.  On  the  dualistic  hypothesis,  two 
varieties  of  chromidia  occur,  one  originating  from  the  vegetative  nucleus  (macro- 
nucleus),  being  chromidia  in  the  restricted  sense,  the  other  derived  from  the 
reproductive  or  micronucleus  being  termed  sporetia.  Chromidia  consist  of  altered 
(?  katabolic)  nuclear  material. 


CLASSIFICATION   OF   THE    PROTOZOA  27 

The  nucleus  plays  the  same  part  in  the  life  of  the  single  celled  organisms  as  it 
does  in  the  cells  of  the  Metazoa  and  Metaphyta.  It  appears  to  influence  in  a  certain 
manner  all,  or  at  least  most,  of  the  processes  of  life,  such  as  motility,  regeneration, 
growth,  and  generally  also  digestion.  Its  principal  influence,  however,  is  exercised 
in  the  propagation  of  the  cells,  as  this  is  always  brought  about  by  the  nucleus. 

The  PROPAGATION  of  the  Protozoa  is  effected  either  by  division  or  by  means  of 
direct  budding.  In  division,  which  is  preceded  by  direct  or  indirect  (mitotic) 
division  of  the  nucleus,  the  body  separates  into  two,  several,  or  even  a  great  many 
segments.  In  this  process  the  entire  substance  of  the  body  is  involved,  or  a  small 
residual  fragment  may  be  left,  which  does  not  undergo  further  division  and  finally 
perishes.  In  the  budding  method  of  multiplication  a  large  number  of  buds  are 
formed,  either  on  the  surface  or  in  the  interior  of  the  organism.  Where  divisions  or 
buddings  follow  one  another  rapidly,  without  the  segments  separating  immediately 
after  their  production,  numerous  forms  develop,  which  are  often  unlike  the  parental 
forms,  and  these  are  termed  swarm  spores  or  spores.  Divisions  imperfectly 
accomplished  lead  to  the  formation  of  protozoal  colonies. 

Sometimes  encystment^  takes  place  previous  to  division.  Frequently,  also, 
sexual  processes  appear,  such  as  the  union  of  two  similar  (isogamous)  or  dissimilar 
(anisogamous)  individuals.  In  the  latter  case  sexual  dimorphism  occurs,  with  the 
formation  of  males  (microgametes)  and  of  females  (macrogametes).  The  union  may 
be  permanent  (copulation),  the  process  being  comparable  with  the  fertilisation  of  the 
ovum  by  a  spermatozoon.  On  the  other  hand,  attachment  may  be  transient  (con- 
jugation) when,  after  the  exchange  of  portions  of  the  nucleus,  the  couple  separate, 
to  multiply  independently  of  each  other.  Sometimes  there  is  an  ALTERNATION  OF 
GENERATIONS,  as  there  may  be  several  methods  of  propagation  combined  in  the 
same  species,  either  direct  multiplication,  conjugation,  or  copulation  being  practised  ; 
the  different  generations  may  thus,  in  certain  cases,  be  unlike  morphologically. 

Protozoa  inhabit  salt  water  as  well  as  fresh  water  ;  they  are  also  found  on  land 
in  very  damp  places,  and  invade  animals  as  parasites. 

Classification  of  the  Protozoa. 

Class  I. — Sarcodina  {Rkizopoda).  Protozoa,  the  body  substance  of  which  forms 
pseudopodia  ;  many  of  them  are  capable  of  developing  chitinous,  chalky,  or  siliceous 
coverings  or  skeletal  structures,  which,  however,  permit  the  protrusion  of  the 
pseudopodia  either  over  the  entire  periphery  or  at  certain  points.  They  possess  one 
nucleus  or  several. 

Order  i. — Amoebina  (Lobosa)  naked  or  with  a  simple  shell,  sometimes  formed 
of  a  foreign  substance  ;  the  pseudopodia  may  be  lobose  or  finger-shaped  ; 
there  may  be  a  contractile  vacuole  ;  generally  only  one  nucleus.  They 
live  in  fresh  or  salt  water,  in  the  soil,  and  also  parasitically. 
Order  2. — Foraininifera  (Reticulata).  Mostly  provided  with  a  calcareous 
shell,  usually  consisting  of  several  chambers,  and  allowing  the  protrusion 
of  the  pseudopodia  either  at  the  periphery  or  only  at  the  opening.  The 
pseudopodia  are  filamentous  and  frequently  anastomosed  ;  there  is  no 
contractile  vacuole  ;  there  are  usually  several  nuclei.  Mostly  marine. 
Order '^i. — Heliozoa.  Naked,  or  with  a  chitinous  or  simple  radial  siliceous 
skeleton  ;  the  pseudopodia  are  filamentous,  and  are  frequently  supported 

'  Independently  of  propagation,  many  protozoa  protect  themselves  from  death  by  encyst- 
ment  when  the  water  in  which  they  are  living  dries  up  ;  in  this  condition  the  wind  may  carry 
them  over  wide  tracts  of  land. 


28  THE   ANIMAL   PARASITES   OF   MAN 

by   firmer  axes,  which    exhibit  no  tendency  to  anastomosis  ;    there  is  a 
contractile  vacuole  ;   one  or  several  nuclei.     Live  in  fresh  water. 
Order  ^.—Radiolaria.      The  body  has  radially-disposed  filamentous  pseudo- 
podia,  and  the  nucleus  is  hidden  in  the  central  capsule  ;  there  is  almost 
always  a  siliceous    framework,    consisting   of   pieces    arranged   radially, 
tangentially,  or  lattice-like  ;    there  is  no  contractile  vacuole,  but   fluid- 
containing  hydrostatic  vacuoles  are  present  in  the  peripheral  protoplasm. 
Marine. 
Class  II. — Mastigophora    {Flagellata).      Protozoa   with    one    or    several    long 
flagella  used  for  locomotion  and  for  acquiring  food ;    in  stationary  forms  their  only 
function  is  to  take  in  food.       Cvtostome  and  contractile  vacuole  mav  be    present. 
Maybe  either  naked  or  provided  with  protective  coverings;    one  or  more  nuclei. 
They  live  either  in  fresh  or  salt  water,  or  may  be  parasitic. 

This  class  is  again  divided  into  several  sub-classes  and  orders,  of  which  only  the 
Euflag>:llata,  with  the  Protomonadina  and  Polymastigoda  are  of  interest  here. 

Class  III. — Sporozoa.     Protozoa  that  only  live  parasitically  in  the  cells,  tissues, 
or  organs  of  other  animals.     They  ingest  liquid  food  by  osmosis  ;  the  surface  of  the 
body  is  covered  with  an  ectoplasmic  layer,  or  cuticle  ;  they  have  no  cilia  in  the  adult 
state,  but  may  form  pseudopodia.     Flagella  occur,  but  only  on  the  male  propagating 
individuals.     There   may  be  one  or    numerous    nuclei,  but   no    contractile  vacuole. 
Propagation  by  means  of  spores,  mostly  provided  with  sporocysts,  is  characteristic. 
Sub-class  I. — Telosporidia.    These  are  usually  of  constant  form,  rarely  amoeboid  ; 
they  are  uninucleate  in  the  mature  state;  they  live  within  host  cells  in  the 
first  stage.     Spore-formation  occurs  at  the  end  of  the  life-cycle. 
Order  i. — Gregari?iida.    Body  of  a  constant,  usually  elongate  form,  surrounded 
by  a  cuticle,     In  the  early  stage  they  lead  an  intracellular  existence;  in 
the  mature  stage  they  live  within  the  intestine  or  body  cavity  of  inverte- 
brate animals,  especially  the  Arthropoda,  and,  like    intestinal   parasites, 
are   provided    with    clinging    organs.       Copulation    usually    isogamous  ; 
the   spores   have   coats    (chlamydospores)    and    usually   contain    several 
minute  germs  (sporozoites). 
Order  2. — Coccidiidea.     Body  of  uniform   spherical  or  oval  shape  :  they  lead 
an  intracellular  life,  but  are  not  freely  motile    in    cavities    of  the  body. 
Fertilization  is  anisogamous  ;  the  spores  have  coals  or  shells  (sporocysts), 
and  usually  contain  several  sporozoites.    Exhibit  alternation  of  generations. 
Order  3. — Hcemosporidia.     Parasites  of  the  blood  corpuscles   of  vertebrate 
animals  ;  they  exhibit  amoeboid  movement;  fertilization  is  anisogamous; 
many  present  alternation  of  generations  and  hosts ;  spores  naked. 
Sub-class  2. — Neosporidia.     They  are  multinucleate  when  adult,  and  the  form  of 
the   body  varies    exceedingly    (often    amoeboid)  ;    spore-formation    com- 
mences before  the  completion  of  growth. 
Order  i. — Myxosporidia.     The  spores  have  valvular  coats,   with  or  without 
caudal  appendages,  with  two,  rarely  four,  polar  capsules.     They  live  free 
in  such  organs  as  the  gall  or  urinary  bladder,  but  are  chiefly  found  in 
connective  tissue.     They  occur  especially  in  fishes. 
Order    2. — Microsporidia.      Spores   with    coats    or   sporocysts ;    no    caudal 
appendage,  with  one  polar  capsule.     They  usually  live  in  the  tissues  of 
Arthropoda. 
Order  3. — Sarcosporidia.      Elongate    parasites    of    the   muscular    fibres    of 
amniotic  vertebrates,  on  rare  occasions  they  occur  also  in  the  connective 
tissue  ;   the  spores,  which  are  kidney  or  sickle-shaped,  are  naked  and 
apparently  have  no  obvious  polar  capsule. 


SARCODINA 


29 


Order  ^. — Haplosporidia.  Simple  organisms,  forming  simple  spores;  they 
occur  in  Rotifers,  Polychaetes,  Fish  and  Man. 

Class  IV. — Infusoria  {Ciliata).  The  body  is  generally  uniform  in  shape,  with 
cilia  and  contractile  vacuole,  frequently  also  with  cytostome  ;  usually  has  macro- 
and  micro-nucleus  ;  live  free  in  water  and  also  parasitically. 

The  "orders  Holotricha.,  Heterotricha,  Olicrotricha^  Hypotricha  and  Peritricha 
are  classified  according  to  the  arrangement  of  the  cilia. 

Class  V. — Suctoria.  Bodies  with  suctorial  tubes,  contractile  vacuoles,  macro- 
and  micro-nucleus,  no  cytostome.  They  generally  invade  aquatic  animals  as  cavity 
parasites,  yet  also  attack  plants  ;  early  stage  ciliated.  Live  sometimes  as  parasites 
on  Infusoria.    [The  Suctoria  are  frequently  regarded  as  a  sub-class  of  the  Infusoria.] 

The  Protozoa  and  Protophyta  are  sometimes  united  under  the  term  Protista 
(Haeckel,  i866).     The  Spirochaetes  are  Protists  (see  pp.  1 14 — 128). 


Class  I.     SARCODINA,  Butschli,  1882. 
Order.     Amoebina,  Ehrenberg. 
A.    Human  Intestinal  Amoebse. 

The  first  record  of  the  occurrence  of  amceba-like  organisms  in  the  human  intes- 
tine, that  is,  in  intestinal  evacuations,  was  that  of  Lambl  (1859) ;  nevertheless,  the  case 
was  not  quite  conclusive,  as  the  occurrence  of  testaceous  amoebae  of  fresh  water 
{Arcella,  Difftiigid)  was  also  reported.  In  1870  Lewis  found  amoebae  associated  with 
disorders  of  the  large  intestine  in  patients  in  Calcutta.  A  year  later  Cunningham 
reported  from  the  same  locality  that  he  had  observed  on  eighteen  occasions,  in  one 
hundred  examinations  of  dejecta  from  cholera  patients,  colourless  bodies  with  amoeboid 


Fig.   I. — Avuxba  co/i,  Lcisch,  in  the  intestinal  mucus.     (After  Losch.) 


movements,  which  became  encysted  and  multiplied  by  fission.  The  daughter  forms 
were  said  to  be  capable  of  dividing  again,  but  they  might  also  remain  in  contact. 
Contractile  vacuoles  were  not  noticed.  The  same  bodies  were  observed  also  in 
simple  diarrhoea  (twenty-eight  cases  out  of  one  hundred.) 

The  case  reported  by  Losch  in  1875  attracted  more  attention.     It  was  that  of 
a  peasant,  aged  24,  who  came  from  the  province  of  Archangel.     He  was  admitted 


30  THE   ANIMAL   PARASITES   OF   MAN 

into  Eichwald's  clinic  at  Petrograd  with  symptoms  of  dysentery.  In  the  dis- 
charges containing  blood  and  pus,  Losch  found  amcebae  in  large  numbers.  When 
at  rest  these  amoebae  measured  from  20 /i  to  35  ft ;  in  a  state  of  movement  their  length 
might  extend  up  to  60  /i  (fig.  i).  The  pseudopodia  appeared  only  singly,  and, 
since  they  were  hyaline  (ectoplasmic),  were  thus  distinguished  from  the  markedly 
granular  endoplasm  that  enclosed  a  spherical  nucleus  of  from  5  /u  to  7  ;u  in  diameter. 
One  or  more  non-contractile  vacuoles  were  present.  Quinine  enemata  had  the 
effect  of  making  the  amoebae  disappear  from  the  fasces  and  thus  causing  the  diarrhoea 
to  abate.  Four  months  after  admission  the  patient  died  from  the  results  of 
intercurrent  pneumonia.  At  the  autopsy  ulceration  of  the  large  intestine  was  found, 
especially  in  the  lower  parts.  Losch  connected  the  amoebae  with  the  ulcerations  by 
experiments  made  on  four  dogs  by  injecting  them  with  recently  passed  stools 
{per  OS  et  anum).  Eight  days  after  the  last  injection  numerous  amcebae  were  found 
in  the  faeces  of  one  of  these  dogs  ;  eighteen  days  after  the  injection  the  animal 
was  killed.  The  mucosa  of  the  rectum  was  inflamed,  covered  with  blood-stained 
mucus  and  ulcerated  in  three  places.  Numbers  of  amoebae  were  found  both  in 
the  pus  of  the  ulcers  and  in  the  mucus.  The  three  other  dogs  remained  healthy. 
From  these  observations  Losch  concluded  that  the  species  of  amoeba  described  by  him 
as  Amccba  colt  could  not  be  regarded  as  the  primary  cause  of  the  disease,  but  that  it 
was  certainly  capable  of  increasing  a  lesion  of  the  large  intestine  already  present, 
or  at  least  of  preventing  its  healing. 

B.  Grassi  (1879)  found  in  the  stools  of  healthy  as  well  as  in  those  of  diarrhoeic 
patients  from  various  localities  in  Northern  Italy,  amoebae  similar  to  those  discovered 
by  Losch.  As  this  was  of  frequent  occurrence,  the  pathogenicity  could  not  be 
definitely  established.  Normand,  formerly  naval  surgeon  at  Hong-Kong,  observed 
numerous  amoebae  in  the  dejecta  of  two  patients  suffering  from  colitis. 

Many  further  investigations,  which  cannot  be  quoted  in  detail,  showed  not  only 
that  intestinal  amoebce  were  widely  distributed  in  man,  but  indicated  with  greater 
certainty  their  role  as  agents  of  dysentery.  The  Commission  sent  out  by  the  German 
Government  in  the  year  1883  to  investigate  cholera  in  India  and  Egypt — whose 
members  discovered  the  cholera  bacillus — also  collected  information  with  regard  to 
dysentery.  In  five  cases  of  dysentery  examined  posi  mortem  at  Alexandria,  with 
the  exception  of  one  case  in  which  ulceration  of  the  colon  had  already  cicatrized  or 
was  approaching  cicatrization,  R.  Koch  found  amoebae  as  well  as  bacteria  in  sections 
from  the  base  of  the  ulcers,  although  such  had  previously  escaped  notice  in  examina- 
tion of  the  dejecta.  Encouraged  by  these  results,  Kartulis  (1885),  who  had  discovered 
amceba-like  bodies  in  the  stools  of  patients  suffering  from  intestinal  complaints  at 
Alexandria,  continued  his  investigations.  The  results,  obtained  from  more  than  500 
cases,  gave  rise  to  the  theory  that  typical  dysentery  vviis  caused  by  amoebae  as  were 
also  the  liver-abscesses  that  often  accompany  it.  Kartulis  supported  his  theory  not 
only  by  the  regular  occurrence  of  amoebae  in  the  stools  of  dysenteric  patients  and 
their  absence  in  other  diseases,  and  by  the  occurrence  of  the  parasites  in  ulcers  of  the 
large  intestine  and  in  the  pus  from  liver-abscesses,  but  also  by  experiments  which 
he  performed  on  cats.  These  were  infected  by  injection /^r  a«//;//  of  stool  material 
rich  in  amoebae  from  subjects  of  dysentery.  The  infection  took  place  also  when 
amoeba-containing,  but  bacteria-free,  pus  from  liver-abscesses  was  used.  It  has  been 
objected  that  the  infection  of  man  with  Amceba  coli^  as  the  dysenteric  amoebae  were 
then  generally  designated,  does  not  take  place  j2^.?r  attu7n  hnx.  per  os.  This  difficulty, 
however,  diminished  in  proportion  as  the  encysted  states  of  amoebae  (fig.  2),  long 
known  in  the  case  of  other  Protozoa,  became  understood.  The  infection  of  man 
(Calandruccio,  1890)  and  of  cats  (Quincke  and  Koos)  succeeded  solely  when  material 
containing  such  stages  was  used.  Amoebae  introduced  into  the  intestine  multiply 
thereby  fission  (Harris,  1894).    However,  this  theory,  to  which  various  other  authors 


SARCODINA  21 

gave  support  on  the  grounds  of  their  own  observations,  encountered  opposition.  Thus 
it  was  estabbshed  that  amoebas  were  not  found  in  patients  in  every  place  where 
dysentery  was  endemic,  or  else  they  were  much  rarer  than  was  expected.  Further 
amoebae  were  present  in  the  most  varied  kinds  of  intestinal  diseases,  both  of  infective 
and  non -infective  characters.     Also  they  were  present  in  quite  healthy  persons. 

Moreover,  for  various  reasons,  infection  experiments  on  animals  failed  to  supply 
proof,  and  finally  a  bacterium  was  discovered  (Shiga,  1898)  to  be  the  excitant  of  one 
form  of  dysentery.  Agglutination  attested  the  specific  part  played  by  this  organism, 
as  it  was  produced  by  the  blood  serum  of  a  person  suffering  from  or  recovered  from 
dysentery,  but  not  by  the  serum  of  one  who  was  uninfected.  Bacillary  dysentery 
consequently  was  a  distinct  entity.  The  final  step  to  be  taken  was  to  decide  whether 
there  was  a  specific  amoebic  enteritis  (amoebic  dysentery  or  amoebiasis,  according  to 
Musgrave). 


Fig.  2. — Encysted  intestinal  amoebae  showing  nuclear  muliiplication. 
(Afier  B.  Grassi.) 

This  question  should  decidedly  be  regarded  from  the  positive  point  of  view. 
It  is  intimately  connected  with  another,  namely,  whether  there  are  not  several  species 
of  intestinal  amoebae.  The  possibility  of  this  had  already  been  recognized.  In 
addition  to  the  Amceba  coli  Losch,  R.  Blanchard  distinguished  yet  another,  Amoeba 
intesiinalis^  and  designated  thereby  the  large  amoebae  described  in  the  first  com- 
munication made  by  Kartulis  ;  later  on  he  stated  the  distinction  between  the  species. 
Councilman  and  Lafleur  ^  (1891)  considered  the  amceba  of  dysentery  to  h&  Amceba  coli 
Losch  and  so  re-named  the  species  Amoeba  dysentericd.  Kruse  and  Pasquale  (1893) 
employed  the  same  nomenclature,  but  retained  the  old  name  Amoeba  coli  Losch  for 
the  non-infectious  species.  Quincke  and  Roos  (1893)  set  forth  three  species:  a 
smaller  species  (25  /x)  finely  granular,  pathogenic  for  men  and  cats  {Ajnosba  coli 
Losch)  ;  a  larger  species  (40  /*)  coarsely  granular,  pathogenic  for  men  but  not  for 
cats  {A.  coli  mitis)  ;  and  a  similar  species  non-pathogenic  either  for  man  or  cat 
{A.  intestini vulgaris),   Celli  and  Fiocca  (1894-6)  went  still  further,  they  distinguished  : 

(i)  Amoeba  lobosa  vdiueiy gutlula  {=A.guttula  Duj),  variety  oblonga  (=A.  oblonga 
Schm.)  and  variety  coli  (=  A.  coli  Losch). 

(2)  Amoeba  spinosa  n.  sp.  occurring  in  the  vagina  as  well  as  in  the  intestine  of 
human  patients  suffering  from  diarrhoea  and  dysentery. 

(3)  Amoeba  diaphana  n.  sp.  found  in  the  human  intestine  in  cases  of  dysentery. 

(4)  Amoeba  vermicularis  Weisse,  present  in  the  vagina  and  in  dysentery  ;  and 

(5)  Amoeba  reticularis  n.  sp.  in  dysentery. 

Shiga  distinguished  two  species;  a  larger  pathogenic  species  with  a  somewhat 
active  movement,  and  a  small  harmless  species  with  a  somewhat  sluggish  movement. 
Bowman  mentions  two  varieties,  Strong  and  Musgrave  (1900)  two  species— the  patho- 
genic Amceba  dysenteries  and  the  non-pathogenic  Amoeba  coli ;  Jager  (1902)  and 
Jiirgens  (1902)  mention  at  least  two  species.  In  the  following  year  (1903)  a  work  by 
Schaudinn  was  published  which  marked  a  real  advance.  This,  in  conjunction  with 
the  establishing  of  a  special  genus  {^Endamoeba  or  Entainceba)  for  human  intestinal 
amoebae  first  by  Leidy^  and  then  by  Casagrandi  and  Barbagallo,^  for  the  time  cleared 
up  the  confused  nomenclature,  the  old  name  Amoeba  coli  being  retained  for  the 

'  '•  Amcebic  Dysentery,"  Johns  Hopkins  Hosp.  Repts.,  ii,  pp.  395-548,  7  plates. 
-  "On  Amceba  blattae,''  Proc.  Acad.  Nat.  Set.,  Philadelphia  (1879),  xxxi,  p.  204. 
3  '■''  Entamceba  hominis  s.  A?nceba  coli  (Losch).     Annali  d'lgiene  speriment.  (1897),   vii, 
p.  103.     See  also  further  remarks  on  p.  34. 


32 


THE   ANIMAL   PARASITES   OF   MAN 


harmless  intestinal  amoebae  of  man,  whereas  the  pathogenic  species  was  desij^inated 
Entama'ba  histolytica.  The  history  of  more  recent  work  is  incorporated  in  the 
accounts  of  the  entamoebas  given  below. 

Entamceba  coll,   Losch,  1875,  emend.  Schaudinn,   1903. 

Syn.  :  Amceba  coli,  Losch,  1875.     EntauKsba  /wniinis^  Casagr.  et  Barbag.  1897. 

The  amoeboid  trophozoite,  according  to  Losch,  measures  26  yu,  to 
30  fjL  and  upwards;  according  to  Grassi  8  /i  to  22 /x  ;  according  to 
Schuberg  12  /x  to  26  /x.  A  separation  of  the  body  substance  into 
ectoplasm  and  endoplasm  is  only  perceived  during  movement.  The 
pseudopodia,  which  are  generally  only  protruded  singly,   are  broad 


Fig.  3. — Entamceha  coli :  life-cycle,  a  —  e,  stages  in  binary  fission  ;  A — Z>,  schizogony,  wiih 
formation  of  eight  merozoites  ;  2 — 10,  cyst  formation  cr  sporogony,  with  formation  of  eight 
nucleate  cysts.     (After  Castellani  and  Chalmers). 

and  rounded  at  the  end  (lobopodia)  and  are  hyaline,  while  the 
remainder  of  the  body  is  granular.  The  ectoplasm  is  less  refractile 
than  the  rest  of  the  cytoplasm;  it  also  stains  less  intensely  (fig.  i), 
and  is  best  seen  on  protrusion  of  a  pseudopodium.  Red  blood 
corpuscles  are  larely,  if  ever,  found  ingested  in  the  cytoplasm. 

The  nucleus  is  vesicular,  and  is  spherical  when  inactive,  measuring 
5 /A   to  yz-t,  with   a   thick   nuclear  membrane.     In  the  centre  of  the 


ENTAMCEBA   CO  LI  33 

nucleus  is  a  chromatinic  body  or  karyosome  or  sometimes  several 
small  nuclear  bodies  formed  of  plastin  and  chromatin  ;  the  remaining 
chromatin  is  arranged  on  the  achromatic  network  in  the  form  of  fine 
granules,  especially  thickly  deposited  on  the  nuclear  membrane. 

Entamoeba  coll  lives  as  a  commensal  in  the  upper  portion  of  the 
large  intestine,  where  the  faeces  still  possess  a  pulpy .  consistency. 
With  their  concentration  and  change  in  reaction  lower  in  the 
bowel,  the  parasites  either  die  or  else  if  they  are  at  a  suitable  stage  of 
development  form  resistant  cysts.  These  cysts  (fig.  2)  can  be  found 
in  great  abundance  in  normal  faeces,  as  Grassi  first  observed.  Slight 
laxantia  or  intestinal  diseases  of  any  kind  producing  increased 
peristalsis,  however,  show  amoebae  even  in  the  unencysted  condition, 
provided  that  the  person  harbours  intestinal  amoebae  generally.  The 
intensity  of  infection  varies  according  to  the  locality;  thus  Schaudinn 
found  that  50  per  cent,  of  the  persons  examined  w^re  infected  with 
harmless  amoebae  in  East  Prussia,  20  per  cent,  in  Berlin  and  about  66 
per  cent,  on  the  Austrian  littoral. 

The  life-history  (fig.  3)  of  the  parasite  exhibits  two  phases  : 
(a)  asexual  multiplication  in  the  intestine,  either  by  binary  fission  or 
by  schizogony  with  formation  of  eight  merozoites,  and  (b)  sporogony 
leading  to  the  production  of  eight-nucleate  cysts.  Infection  results 
from  ingestion  of  cysts.  Only  cysts  with  eight  nuclei  are  infective. 
The  diameter  of  such  cysts  is  about  15  /^  to  20  fM. 

There  are  varying  accounts  of  the  details  of  the  life-cycle  of  Entamosba  colt  in  its 
different  stages.  Thus,  regarding  schizogony  or  multiple  fission  it  was  formerly 
stated  that  the  nucleus  of  the  parent  amoeba  divided  into  eight  portions,  which  after 
dissolution  of  the  nuclear  membrane,  passed  outwards  into  the  cytoplasm,  which 
segregated  around  each.  Eight  merozoites  were  thus  produced.  More  recently  the 
process  of  schizogony  has  been  considered  to  consist  in  the  repeated  division  of  the 
nucleus  into  two,  four,  and  finally  eight  nuclei  (fig.  3,  A — d),  and  the  formation  of 
eight  merozoites  or  amoebulas. 

The  process  of  encystment  is  initiated  by  the  extrusion  of  all  liquid  and  foreign 
bodies  from  the  protoplasm,  which  assumes  a  spherical  form  (fig.  4,  a).  The  rounded 
uninucleate  amoeba  then  secretes  a  soft  gelatinous  coat,  which  finally  diffisrentiates 
into  a  double  contoured  cyst  wall  in  older  cysts.  According  to  Casagrandi  and 
Barbagallo,  the  size  of  the  cyst  varies  from  8  ju  to  30  /*,  and  averages  about  15  /*. 
According  to  Schaudinn  (1903)  the  cytological  changes  during  cyst  formation  are  as 
follows.  The  nucleus  of  a  rounded  uninucleate  form  divides  into  two  (fig.  4,  B). 
Each  of  these  nuclei  fragments  into  chromidia  (fig.  4,  C),  some  of  which  are  absorbed, 
while  others  reunite  so  that  the  cell  becomes  binucleate  again.  Each  of  these 
nuclei,  by  a  twice  repeated  division,  produces  three  nuclei  (fig.  4,  D),  the  smaller 
two  of  which  degenerate  and  were  regarded  as  reduction  nuclei.  There  is  a  clear 
zone  or  vacuole  in  the  middle  of  the  cyst  during  these  maturation  processes,  dividing 
the  cyst  into  two  halves.  After  the  nuclear  reduction  the  clear  space  disappears, 
and  each  nucleus  (termed  by  some  a  gamete  nucleus)  divides  into  two  pronuclei 
(fig.  4,  e).  The  pronuclei  of  the  pairs  were  said  by  Schaudinn  to  differ  slightly. 
Copulation  occurs  between  pairs  of  unlike  pronuclei,  and  is  an  example  of  autogamy 
(fig.  4,  F).     When  complete,  each  of  the  fusion  nuclei  (synkarya)  divides  twice,  giving 

3 


34 


THE   ANIMAL   PARASITES   OF   MAN 


rise  first  to  four  and  finally  to  eight  nuclei.  Eight  amcebulae  are  thus  formed  within 
the  cyst. 

According  to  Hartmann  and  Whitmore  (191  OS  however,  autogamy  does  not 
occur  within  the  cysts  of  E.  colt.  They  consider  that  eight  small  amoebulas  are 
formed  (fig.  3,  2—10)  which  escape  from  the  cyst  and  then  conjugate  in  pairs 
(fig.  3,  10 — 12),  afterwards  growing  into  a  new  generation  of  trophozoites. 

Only  some  10  to  20  per  cent,  of  the  cysts  evacuated  with  the  fasces  undergo  the 
full  course  of  development,  the  majority  perish  previously.  In  old  dry  faeces,  only 
cysts  with  eight  nuclei  are  found,  and  it  is  these  alone  that  cause  the  infection. 

Entamceba  ivilliamsi,  E.  biitschlii,  E.  hartmanni  and  E.  poleki  (Prowazek)  are 
probably  only  varieties  of  E.  colt. 


Fig.  4. — So-called  autogamy  of  ^«/a?/za'/^a  ^(7//.  A,  rounded  amoeba;  B,  nucleus  dividing ; 
c,  the  two  daughter-nuclei  giving  off  chromidia ;  d,  each  nucleus  has  formed  two  reduction 
nuclei ;  e,  cyst  membrane  formed,  and  gamete  nuclei  are  dividing ;  f,  cyst  with  two  synkarya. 

The  principal  feature  distinguishing  Entamoeba  coll  from  E, 
histolytica  is  the  formation  of  eight-nucleate  cysts  by  the  former  aS' 
contrasted  with  the  tetra-nucleate  cysts  of  the  latter.  The  cyst-wall  of 
E.  coll  is  thicker  than  that  of  E.  histolytica  (tettagena).  Further,  E.  coli 
does  not  usually  ingest  red  blood  corpuscles,  nor  are  "  chromidial 
blocks"  present  inside  its  cyst  (see  p.  40). 

According  to  Chatton  and  Lalung-Bonnaire'^  (191 2)  the  entamcebae 
of  vertebrates  should  be  placed  in  a  separate  genus  Loschia,  as  they 
differ  in  their  life-history  from  E.  blatta%  the  type  species  of  Entamoeba. 
Leidy  (1879),  however,  named  the  genus  Eudanioeba,  but  further 
researches  are  necessary  on  biological  variation  among  these 
organisms. 

Entamoeba  histolytica,  Schaudinn,  1903. 

Syn.  :  Amoeba  coli,  autt.  p.  p.     Ainosba  dysenfericp,  autt.  p.  p. 

The  average   size  of  the  amoeboid  trophozoite  is  25  fji,  to  30  fi. 

In  fccces  diluted  with  salt  solution  the  amoebae  swell  to  40  ^  and  more. 

There  is  sometimes  separation  of  the  body  substance  into  a  strongly 

refractile  vitreous  ectoplasm  and  a  corneous  endoplasm,  pronounced 


'  Archiv  f.  Protistenktinde^  xxiv,  p.  182. 
-  Bull.  Soc.  Path.  Exotique^  v,  p.  135. 


ENTAMCEBA    HISTOLYTICA 


35 


even  in  repose,  although  the  former  is  not  equally  thick  at  all  parts  of 
the  periphery.  In  the  endoplasm  generally  there  are  numerous  foreign 
bodies  (bacteria,  epithelial  cells,  colourless  and  red  blood  corpuscles 
(fig.  6),  and  occasionally  living  flagellates  of  the  intestine).  The 
nucleus  is  4  /z  to  6  /a  in  diameter,  and  may  be  difficult  to  recognize 
because  it  is  sometimes  weakly  refractile  and  poor  in  chromatin.  Its 
shape  is  slightly  variable  ;  it  is  usually  excentric,  sometimes  wholly 
peripheral  at  the  limit  of  the  two  parts  of  the  body.  Vacuoles  are  not 
present  in  quite  fresh  specimens,  but  appear  later.  In  the  study  of 
E.  Jiisfolytica,  the  morphological  characters  of  the  trophozoite  or 
vegetative  stage  of  the  organism  formerly  separated  as  E.  tetragena 
(figs.  5,  6,  Sa)  must  be  considered  (see  p.  38). 


Fig.  ^.—Entamceba  histolyiica  [^tetragena  form),  showing  three  successive  changes  of  form 
due  to  movement.      X   IICX).      (After  Hartmann.) 


The   history  of  the  development    ot    these   species,  which  give  rise  to  amoebic 
enteritis  as  distinguished  from  bacillary  dysentery,  was  formerly  not  so  well  known 
.as  that  of  E.  coli.     Upon  being  introduced  into  cats  {per  anum)  dysenteric  amoebae 
provoke  symptoms  similar  to  those  in  man.    In  the 
latter,  besides  metastatic  liver  abscesses,  abscesses 
of  the  lungs,  and,  according  lo  Kartulis,  cerebral 
abscesses  are  occasionally  produced.     Marchoux 
(1899)  states  that  when  the  disease  has  lasted  for 
some  time  liver  abscesses  are  produced  in  cats  also. 

In  the  large  intestine  of  infected  cats  the 
amoebae  creep  over  the  epithelium,  and  here  and 
there  they  force  the  epithelial  cells  apart,  as  well 
as  removing  them  or  pushing  them  in  front  of 
them  ;  the  amoebae  thus  insert  themselves  into  the 
narrowest  fissures.  They  penetrate  also  into  the 
glands  through  the  epithelium,  and  thence  into 
the  connective  tissue  of  the  mucosa.  Intestinal 
and  glandular  epithelia  perish  under  the  influence 
of  these  parasites  :  the  cells  are  pushed  aside, 
fall  to  pieces  or  are  absorbed  by  the  amoebae, 
mucosa  the  amoebae  migrate  further,  and  often  accumulate  above  the  muscles. 
Finally  they   rupture    this    and    force    their   way    into    the    submucosa.      In   cats. 


Fig.  6. — Entamaba  histolytica 
which  has  ingested  many  red  blood 
corpuscles.  x  iioo.  (After  Hart- 
mann.) 

In   the   connective  tissue  of  the 


36 


THE   ANIMAL   PARASITES   OF   MAN 


apparently,  the  penetration  is  not  so  great  as  in  men,  according  to  Kruse  and 
Pasquale.  During  their  migration  the  parasites  also  gain  access  to  the  lymph- 
follicles  of  the  wall  of  the  intestine,  which  become  swollen  and  commence 
to  suppurate  ;  follicular  abscesses  arise  and  after  their  rupture  follicular  ulcers. 
The  diseased  patches  in  the  mucosa  are  markedly  hyperaemic  and  numerous  hiemor- 
rhages  are  set  up.  Roos  and  Harris  state  that  the  amoebae  also  oenetrate  into 
the  blood-vessels  (fig.  7)  and  this  explains  the  occurrence  of  metastatic  abscesses.' 
The  whole  submucosa  is  severely  swollen  at  the  diseased  spot  and  undergoes  small- 
celled  infiltration  in  the  neighbourhood  of  the  colonies  of  amoebae.  From  these 
findings  Jiirgens  (1902)  draws  the  conclusion^  which  is  followed  here,  that  the  amoebae 


Fig.  7.— Section  through  wall  of  large  intestine  (of  a  man)  close  under  an  ulcer  caused  by 
Entamoeba  histolytica.  A,  amoebse  that  have  penetrated  partly  in  blood-vessels  (Bv),  partly 
ill  tissue  of  submucosa  to  the  muscularis.      Magnified.     (After  Harris.) 

are  causative  agents  of  the  enteritis  of  cats,  which  disease  is  well  defined,  both 
pathologically  and  anatomically.  Subsequent  researches  confirm  the  experience  of 
earlier  authors  ;  great  precautions  were  taken  to  exclude  errors,  hence,  as  with 
Gross  and  Harris,  no  exception  can  be  taken  to  their  results.  The  inoculation 
material  was  derived  from  soldiers  who  suffered  from  amoebic  enteritis  in  China 
and   who   were  admitted    into  the   garrison   hospital    at    Berlin.     In    order   to    be 


'  Lung  abscesses  generally  arise  by  the  bursting  of  a  liver  abscess  through  the  diaphragm 
into  the  right  lower  lobe  of  the  lung,  sometimes  also  through  conveyance  of  amoebcE  by  means 
of  the  blood-stream  (Banting). 

^  These  findings  were  confirmed  by  Schaudinn  by  means  of  investigations  on  cats  and  men. 
Cf.  also  Alfred  Gross,  Marchoux,  P.  G.  Woolley,  W.  E.  Musgrave,  H.  F.  Harris  and  others. 


ENTAMOEBA   HISTOLYTICA  yj 

independent  of  the  patients  themselves,  transmission  experiments  from  cat  to  cat 
were  performed,  after  the  first  experiments  on  cats  yielded  positive  results.  This 
was  also  effected  by  rectal  feeding  as  employed  by  earlier  workers.  Such  appeared 
necessary  in  order  to  prevent  the  evacuation  of  the  inoculation  material /^r  «««w, 
as  well  as  to  avoid  the  employment  of  morphia  and  ether  narcosis.  Forty-six 
cats  were  used  for  the  experiments.  Ten  cats  received  tested  stools  containing 
motile  amoebie  from  soldiers  suffering  from  amoebic  enteritis  contracted  in  China. 
Sixteen  other  cats  received  stools  from  cats  infected  by  inoculation.  All  the  animals 
sickened  and  suffered  from  the  disease.  Five  cats  received  dejecta  from  human 
amoebic  enteritis  in  which,  however,  no  motile  amcebae  were  present.  Thirteen  cats 
received  stools  from  soldiers  who  suffered  from  bacillary  dysentery.  None  of  the 
latter  cats  took  the  complaint  and  none  showed  changes  in  the  large  intestine  upon 
sectioning.  The  injection  of  various  bacteria,  obtained  from  a  stool  of  amoebic 
enteritis  pathogenic  to  cats,  remained  without  result  in  both  the  cats  employed  for 
this  experiment.  Lastly,  two  cats,  which  had  been  kept  with  those  artificially 
infected,  were  taken  ill  spontaneously  and  suffered  from  the  disease.  In  the  opinion 
of  Harris,  who  ascertained  the  harmless  nature  of  bacteria  derived  from  the 
intestinal  flora  containing  dysenteric  amoebae,  young  dogs  are  capable  of  being 
infected. 

Within  the  large  intestine  an  active  increase  of  Entamoeba  histolytica  must  occur. 
Nevertheless,  Jiirgens  did  not  definitely  find  changes  that  might  be  interpreted  in  this 
sense.  Schaudinn  (1903)  observed  division  and  gemmation  in  vivo.  Both  processes, 
in  which  the  nucleus  divides  by  amitosis,  can  only  be  distinguished  by  the  fact  that 
the  daughter  individuals  are  similar  in  binary  fission  but  dissimilar  in  gemmation, 
whether  they  make  their  appearance  singly  or  in  greater  numVjers.  Schizogony, 
resulting  in  the  formation  of  eight  individuals,  which  is  so  characteristic  for  Ent- 
amoeba coli,  was  not  observed.  (But  schizogony,  into  four  merozoites,  is  now  known 
to  occur.     Gemmation  processes  are  apparently  degenerative.) 

Resistant  stages,  which  serve  for  transmission  to  other  hosts,  are  according  to 
Schaudinn'  first  formed  when  the  diseased  portions  commence  to  heal,  or  more 
accurately,  the  recovery  commences  when  the  vegetative  increase  of  the  amoebae  in  the 
intestine  discontinues.  The  so-called  spores  of  E.  histolytica  were  distinguished  very 
definitely  from  those  of  E.  coli;  they  were  said  to  consist  of  spheres  of  only  3  to 
7  /i  in  diameter,  which  were  surrounded  by  a  double  membrane,  at  first  colourless,  but 
becoming  a  light  brownish  yellow  colour  after  a  few  hours,  and  possessing  a  protoplas- 
mic content  containing  chromidia.  They  were  said  to  arise  by  fragments  of  chromatin 
passing  outwards  from  the  nucleus  of  the  amoeba  into  the  surrounding  cytoplasm 
(fig.  9,  a)  and  undergoing  so  marked  an  increase  that  finally  the  whole  cytoplasm 
became  filled  with  chromidia.  The  remainder  of  the  nucleus  underwent  degenera- 
tion and  became  extruded.  On  the  surface  of  the  cytoplasm  there  then  arose  small 
protuberances  containing  chromidia.  These  processes  had  been  observed  in  the 
living  organisms.  They  gradually  divided  and  separated  from  membranes  which  later 
became  yellow.  The  remainder  of  the  amoeba  perished.  Craig^  had  also  seen  phases 
of  this  process  of  development.  It  must  be  remarked  that,  according  to  recent 
researches,  these  processes  of  exogenous  sporulation  are  degenerative  in  character  (see 
p.  41).  The  small  spores  may  be  fungi.  The  "  sporulation  "  processes  are  only  men- 
tioned here  as  a  warning.  They  are  now  only  of  historic  interest.  By  means  of  an 
experiment  made  on  a  cat,  Schaudinn  ascertained  that  ingestion  of  permanent  cysts, 
which  resist  desiccation,  is  the  cause  of  the  infection.  The  animal  took  food  containing 
dry  faeces  with  amoeba  cysts  ;  these  fasces  came  from  a  patient  suffering  from  amoebic 

'  Arb.  a.  d.  kaiserl.  Gesundheitsamtey  xix,  pp.  547-576. 

2  "  Life  cycle  of  Amoeba  coli  in  Human  Body,"  American  Medicine,  1904,  vii,  p.  299  ;  viii, 
p.  185. 


58  THE   ANIMAL   PARASITES   OF   MAN 

enteritis  in  China.  On  the  evening  of  the  third  day  the  cat  evacuated  blood-stained 
mucous  faeces  which  contained  large  numbers  of  typical  Entamceba  histolytica.  On 
the  fourth  day  after  the  infection  the  animal  experimented  upon  died,  and  the  large 
intestine  showed  the  changes  previously  stated. 

E  histolytica  also  is  found  in  the  large  intestine.  This  was  originally  shown 
to  be  the  case  by  Kartulis,  and  the  fact  has  recently  been  confirmed  from  many 
quarters.  It  is  also  present  in  the  metastatic  abscesses  of  which  it  is  the  cause 
{cf.  among  other  authors,  Rogers,  Brit.  Med.  Journ.,  1902,  ii,  No.  2,177,  P-  844  ;  and 
1903,  i,  No.  2,214,  P-  1315)- 

It  should  lastly  be  pointed  out  in  this  connection  that  mixed  infections  also  take 
place.  For  instance,  in  addition  to  E.  histolytica^  E.  coli,  and,  under  certain  circum- 
stances, flagellates  may  be  found  together.  In  the  same  way  E.  coli  may  come 
under  observation  even  in  bacillary  dysentery.  On  the  other  hand,  Schaudinn 
stated  that  in  cases  of  dysentery  endemic  in  Istria,  Entamoeba  coli,  if  it  had  hitherto 
been  present,  disappeared,  to  return  again  after  recovery  from  the  illness. 


^■  «s 


Fig.  8. — Entama-ba  histoiy.i.u.  a,  trophozoite  {tetragena  type)  containing  red  blood 
corpuscles,  x  1,300;  h  and  c,  two  isolated  nuclei  showing  different  appearances  of  karyosome, 
centriole  and  nuclear  membrane,  x  2,600.     (After  Hartmann.) 

(Entamceha  tetragenUy  Viereck,  1907.) 

This  amoeba  must  now  be  considered  to  be  a  part  of  the  hfe- 
cycle  of  Entamoeba  histolytica,  in  fact  a  very  important  part  of  that 
cycle,  especially  in  its  tetranucleate  cystic  stages. 

This  organism,  the  so-called  Entaniccba  tetragena,  may  occur  in 
the  human  intestine  in  cases  of  amoebic  dysentery,  especially  in 
mild  or  chronic  cases.  It  was  discovered  by  Viereck  in  1907  in 
patients  suffering  from  dysentery  contracted  in  Africa.  Soon  after- 
wards an  independent  description  was  published  by  Hartmann,  who 
called  the  ainoeba  E.  africana.  It  was  also  studied  by  Bensen  and 
Werner.  Recently  (1912-13)  much  work  has  been  published  on 
this  amoeba  by  Darling  and  others  ;  in  this  way  its  relationship  to 
Schaudinn's  E.  histolytica  has  been  made  known. 

In  general  morphology  it  somewhat  resembles  Entamceha  coli,  and 


ENTAMOEBA   HISTOLYTICA    (TETRAGENA) 


39 


its  discoverer  at  first  mistook  it  for  a  variety  of  that  species.  Accord- 
ing to  Hartmann,  a  distinct  ectoplasm  is  only  clearly  visible  when 
a  pseudopodium  is  protruded  (fig.  5).  The  granular  endoplasm  may 
contain  ingested  red  blood  corpuscles  (fig.  6).  The  large,  round 
nucleus  is  visible  in  the  fresh  state  (fig.  8,  a).  So-called  chromidial 
masses  (?  crystalloidal  substances)  may  occur  in  the  cytoplasm. 

Some  investigators,  as  Hartmann,^  lay  stress  on  the  internal 
structure  of  the  nucleus  (fig.  8,  b,  c),  best  seen  in  preparations  fixed  wet 
and  stained  with  iron-haematoxylin.  The  nucleus  is  limited  by  a  well- 
marked  nuclear  membrane,  on  the  inside  of  which  granules  or  nodules 
of  chromatin  may  occur.     There  is  a  karyosome,  which,  in  success- 


FiG.  9. — Entamceba  histolytica  {tetragena  form).  «,  emission  of  chromatin  from  nucleus  ; 
b,  nuclear  division  ;  <r,  degenerating  form  with  two  nuclei  ;  d,  e,  f,  cysts  containing  one,  two 
and  four  nuclei  respectively,  and  showing  chromidial  blocks.     X  2,000.     (After  Hartmann.) 

fully  stained  specimens,  shows,  at  times,  a  central  dot  called  a 
centriole.  (The  nucleus  of  Entamoeba  coll  does  not  contain  such  a 
centriole.)  However,  the  structure  of  the  nucleus  varies  at  different 
periods  during  the  life-cycle. 

The  diameter  of  the  trophozoites  or  vegetative  forms  (fig.  8,  a)  is 
variously  given  as  from  20  jx  to  40  ix.  Multiplication  proceeds  by 
binary  fission  and  also  by  schizogony  into  four  merozoites.^ 

Reproduction  takes  place  by  endogenous  encystment  (fig.  9,  <:/-/), 
which  is  preceded  by  nuclear  division  into  two,  reduction  and  then 
autogamy.     The  interpretation  of  the  latter  phenomenon  as  autogamy 


'  Arch.f.  Protistenkunde  (1911),  xxiv,  p.  163. 

-  See  Darling,  1913,  Arch.  Inteni.  Med.,  vol.  ii,  pi.  i,  fig.  3- 


40  THE   ANIMAL   PARASITES   OF   MAN 

is  disputed  by  some  authors.  The  round  cysts,  which  may  measure 
i2yL6  to  15 /z,  in  diameter,  contain  four  nuclei,  together  with  darkly 
staining  masses  of  various  shapes,  the  so-called  '*  chromidial  blocks" 
(fig.  9,  /).  The  cyst-wall  of  E.  histolytica  (tetragena)  is  thinner  than 
that  of  E.  coli,  and  the  diameter  of  the  cyst  is  rather  less.  E.  histolytica 
has  not  yet  been  cultivated. 

Infection  in  man  occurs  by  way  of  the  mouth  by  the  ingestion 
of  cysts.  A  patient  showing  acute  symptoms  of  dysentery  is  not 
usually  infective,  for  he  is  merely  harbouring  the  large  trophozoites, 
which,  by  experiment,  have  been  shown  not  to  be  infective  to  animals 
(kittens)  when  administered  by  the  mouth.  The  stools  of  recovered 
patients  may  still  contain  cysts,  and  they  may  thus  act  as  cyst-carriers 
or  reservoirs  of  disease  by  infecting  w^ater  and  soil.  The  stools  of  such 
cyst- carriers  are  often  solid,  and  so  cysts  of  E.  histolytica  (tetragena) 
are  easily  overlooked.  Mathis  (1913)^  points  out  that  healthy  carriers 
of  E.  histolytica  may  be  found  ;  8  per  cent,  of  the  natives  of  Tonkin 
examined  by  him  were  healthy  carriers  of  cysts. 

In  return  cases,  or  prolonged  untreated  cases  of  entamoebic 
dysentery,  a  generation  of  smaller  trophozoites  is  associated  with,  or 
replaces  the  larger  ones.  In  stools  they  are  frequently  refractile  and 
consequently^  stain  slowly  intra  vitani.  These  trophozoites  are  the 
'^  smaller,  senile,  or  pre-cyst  generation  "  of  Darling.  This  pre-cyst 
generation  is  characterized  by  the  presence  of  blocks  of  crystalloidal 
substance  in  the  cytoplasm,  and  by  the  possession  of  a  prominent, 
densely  stainable  karyosome.  Darling  believes  this  generation  to 
be  the  same  as  that  described  by  Elmassian  as  Entainceba  ininuta.^ 
Walker,^  Darling,'^  Wenyon'^  and  others  believe  that  Entamoeba 
histolytica^  which  was  only  seen  by  Schaudinn  in  a  single  case,  that  of 
a  Chinaman,  is  really  £.  tetragena.  Darling  states  that  if  the  published 
illustrations  of  E.  histolytica  and  of  E.  tetragena  are  collected  from  the 
literature  and  compared,  it  will  be  seen  that  the  writers  have  been 
calling  E.  histolytica  the  large  trophozoites  seen  in  dysenteric  stools. 
These  large  trophozoites  frequently  display  no  karyosome,  but  they 
can  be  demonstrated  as  E.  tetragena  by  animal  inoculation,  or  by  the 
history  of  the  case.  On  the  other  hand,  the  illustrations  of  £.  tetragena 
show  that  the  authors  have  been  dealing  with  the  small  generation  or 
reduced  forms  (''  E.  minnta  "),  which  are  the  direct  descendants  of  the 
large  trophozoites.  If  kittens  are  inoculated  rectally  with  dysenteric 
material  containing  large  trophozoites,  the  strain  may  be  carried  in 
successive  kittens  for  four  to  six  transfers.     If,   on  the  other  hand, 

'  Bull.  Soc.  Med.  et  Chirurg.  Indo-Chine,  iv,  p.  474, 

-  Centralbl.f.  Balder.^  Orig.,  lii,  p.  335.  ^  Philip.  Journ.  Sc.  (191 1),  B,  vi,  p.  259. 

••  Annals  Trap.  Med.  and Parasitol.  (1913),  vii,  p.  321. 

^  Brit.  Med.  Joitrn.,  Nov.  15,  1913,  p.  1287,  and  loiirn.  Land,  School  Trap.  Med.,  ii,  p.  27. 


NOC'S   ENTAMOEBA  4I 

kittens  are  inoculated  rectally  with  small  trophozoites  of  the  pre-cyst 
generation,  the  transmission  cannot  be  carried  through  more  than  one 
or  two  kittens.  Wenyon  has  succeeded  in  maintaining  E.  tetragena  in 
kittens  for  several  generations. 

In  some  of  the  preparations  from  the  last  remove,  pathological 
forms  of  the  trophozoites  may  be  seen.  These  show  abnormal  forms 
of  budding,  especially  peripherally,  such  as  have  been  described  by 
Schaudinn  and  by  Craig  as  characteristic  oiE.  histolytica.  Schaudinn's 
small  peripheral,  exogenous  buds  and  cysts  are  thus  explained.  Craig 
has  latterly  changed  his  views. 

Further,  Darling  states  that  tetragena  cysts  fed  by  the  mouth  to 
kittens  produce  bowel  lesions  in  which  trophozoites  having  the  char- 
acters of  E.  tetragena,  E.  histolytica  and  E.  nipponica  (Koidzumi)  occur. 

In  view  of  the  work  of  recent  observers,  the  peculiar  exogenous 
encystment  which  Schaudinn  made  characteristic  of  Entamoeba 
histolytica  has  been  shown  to  be  due  to  degenerative  changes  in  senile 
races  of  the  amoeba.  E.  histolytica  and  E.  tetragena  are  one  and  the 
same  species,  and  its  trophozoite  is  subject  to  variation.  According 
to  some  observers  the  histolytica  type  of  nucleus — described  by 
Schaudinn  as  being  poor  in  chromatin  and  not  easily  seen  in  the 
fresh  state — occurs  frequently  in  patients  with  severe  symptoms  of 
dysentery ;  on  the  other  hand,  the  tetragena  type  of  nucleus — round 
and  easily  seen  in  the  fresh  state — may  occur  in  cases  presenting  slight 
dysenteric  symptoms.  Intermediate  t3^pes  of  nuclei  are  seen.  The 
name  of  this  species,  the  principal  pathogenic  amoeba  of  man,  must 
then  be  E.  histolytica  by  priority.  The  cystic  stages  of  E.  histolytica  are 
those  first  recorded  by  Viereck  and  formerly  described  as  E.  tetragena. 
The  geographical  distribution  of  E.  histolytica  is  wide. 

Nog's  Entamoeba  (1909). 

A  species  of  Entamoeba  was  cultivated  by  Noc^  in  1909  from  cysts 
derived  from  liver  abscesses,  from  dysenteric  stools  and  from  the 
water  supply  of  Saigon,  Cochin  China.  He  cultivated  it  in  associa- 
tion with  bacteria.  It  is  pathogenic.  It  has  been  considered  allied 
to  E.  histolytica,  and  shows  internal  segmentation  or  schizogony.  It 
exhibits  polymorphism.  This  amoeba  has  been  found  by  Greig  and 
Wells  (191 1)  in  cases  of  dysentery  in  India.  It  is  an  important 
organism  and  requires  further   investigation. 

Certain  other  Entamcebae^  have  been  described  at  various  times  from 
the  intestinal  tract  of  man.  Probably  most,  if  not  all,  of  these  are  not 
good  species  and  in  some  cases  much  more  information  is  needed. 

Entamoeba  tropicalis  (Lesage,    1908).      This  parasite  is  said  to  be 

'  Noc,  F.  (1909),  Ann.  Inst.  Pasteur^  xxiii.  p.  177. 

-  See  Fantham,  H.  B.  (191 1),  Annals  Trop.  Med.  and ParasitoL,  v,  p.  in. 


42  THE   ANIMAL   PARASITES   OF   MAN 

non-pathogenic,  and  to  occur  in  the  intestine  of  man  in  the  tropics. 
It  has  a  general  resemblance  to  E.  coil,  but  forms  small  cysts  (6  a  to 
lo  fjb  in  diameter).  The  nucleus  of  the  cyst  is  said  to  break  up  into  a 
variable  number  of  daughter  nuclei,  from  three  to  thirteen  having 
been  noted.  Lesage  states  that  it  is  culturable  in  symbiosis  with 
bacteria.     It  is  probably  a  variety  of  E.  coli,  if  not  a  cultural  amoeba. 

Entamoeba  hominis  (Walker,  1908)  has  a  diameter  of  6yLt  to  15//-.  A 
contractile  vacuole  is  present.  Encystment  is  total,  and  small  cysts  are 
formed.  It  is  culturable.  The  original  strain,  now  lost,  was  obtained 
from  an  autopsy  in  Boston  Hospital.  This  organism  is  probably 
a  cultural  amoeba. 

Entamoeha  phagocytoides  (Gauducheau,  1908).  This  parasite  was 
discovered  in  a  case  of  dysentery  at  Hanoi,  Indo-China.  The  amoeba 
is  small,  2  yu,  to  iS/x-  in  diameter.  It  is  active.  It  ingests  bacteria  and 
red  blood  corpuscles,  while  peculiar  spirilla-like  bodies  are  found  in 
its  cytoplasm.  It  multiplies  by  binary  and  multiple  fission.  It  can 
be  cultivated.  More  recently  (191 2)  the  author  appears  to  consider 
the  amoeba  to  be  a  stage  of  a  Trichomonas,  but  abandons  the  view 
later  (19 14).     Further  researches  on  this  organism  are  needed. 

Entamoeba  minuta  (Elmassian,  1909)^  was  found,  in  association 
with  E.  coli,  in  a  case  of  chronic  dysentery  in  Paraguay.  It  resembles 
E.  tetragena  but  is  smaller,  rarely  exceeding  14  //,  in  diameter.  Schizo- 
gony occurs,  four  merozoites  being  produced.  The  encystment  is 
total  and  endogenous,  giving  rise  to  cysts  containing  four  nuclei. 
This  amoeba  is  considered  by  Darling  and  others  to  be  the  pre-cyst 
trophozoite  stage  of  E.  histolytica  (tetragena). 

Entamoeba  nipponica  (Koidzumi,  1909)  was  found  in  the  motions 
of  Japanese  suffering  from  dysentery  or  from  diarrhoea,  in  the  former 
case  in  coriipany  with  Entamceba  histolytica.  Its  diameter  is  15  yu,  to 
30  yu,.  The  endoplasm  is  phagocytic  for  red  blood  corpuscles.  The 
nucleus  is  well  defined,  resembling  that  of  E.  coli  and  of  E.  tetragena. 
Multiplication  occurs  by  binary  fission  and  by  schizogony.  Encyst- 
ment is  total,  but  has  not  been  completely  followed.  Darling  and 
others  consider  that  this  is  an  abnormal  form  of  E.  histolytica, 
while  Akashi  (1913)  doubts  if  it  is  an  amoeba  at  all,  but  rather  is 
to  be  regarded  as  shed  epithehal  cells. 

General  Remark. — It  is  now  considered  by  some  workers  that 
true  Entamoebae  cannot  be  cultivated  on  artificial  media.  Quite 
recently  Williams  and  Calkins  (1913)^  have  somewhat  doubted  this 
opinion,  and  state  that  certain  cultural  amoebae,  originally  obtained 
from  Musgrave  in  Manila,  exhibit  the  various  morphological  variations 
associated  with  true  entamoebae  of  the  human  digestive  tract. 

'  Centralhl.  f.  Bakter.,  Orig.,  lii,  p.  335. 
-  Jotirn.  of  Med.  Research,  xxix,  p.  43. 


ENTAMCEBA   BUCCALIS  43 

Entamoeba  buccalis,  Prowazek,  1904. 

The  size  varies  from  6  /x  to  32  /x.  Ectoplasm  is  always  present ; 
the  endoplasm  contains  numerous  food-vacuoles.  The  nucleus  is 
vesicular,  with  a  greenish  tinted  membrane  which  is  poor  in 
chromatin.  The  size  of  the  nucleus  is  from  1*5 />t  to  4"5yLfc.  A  con- 
tractile vacuole  is  not  visible.  The  pseudopodium  is  broad.  It  was 
discovered  in  the  mouths  of  persons  with  dental  caries  at  Rovigno  and 
also  at  Trieste,  being  most  easily  found  in  dense  masses  of  leucocytes, 
also  among  leptothrix  and  spirochaste  clusters.  It  can  be  easily  dis- 
tinguished from  leucocytes  by  more  intense  staining  with  neutral  red. 
Multiplication  proceeds  by  fission.  Transmission  may  take  place 
through  the  small  spherical  cysts.  This  species  (fig.  10)  has  since  been 
observed  in  Berlin,  and  is  also  occasionally  found  in  carcinoma  of 
various  regions  of  the  oral  cavity.     (Leyden  and  Lowenthal,  1905). 


■■^^ 


Fig.  10. — Entamoeba  buccalis^  Prow,  a-d,  the  same  specimen  observed  during  five  minutes. 
X  1,000.  e,  amoeba  fixed  and  stained  with  iron-hsematoxylin.  x  1,500.  (After  Leyden 
and  Lowenthal.) 

Entamoeba  buccalis,  Prow.,  is  said  to  be  allied  to  a  protozoon  which 
A.  Tietze  has  found  either  encysted  or  free  in  the  lumen  of  the  orifice 
of  the  parotid  gland  of  an  infant  aged  4  months.  The  gland  had 
undergone  pathological  change,  and  had  therefore  been  extirpated. 
The  organisms,  which  were  roundish  and  three  to  four  times  the  size 
of  the  normal  epithelial  cells  of  the  gland,  were  without  a  membrane 
and  possessed  a  nucleus  in  which  the  chromatic  substance  appeared 
to  be  contained  in  a  karyosome.  Bass  and  John's^  (Feb.  191 5)  and 
Smith,  Middleton  and  Barrett  (1914)  state  that  E.  buccalis  is  the  cause 
of  pyorrhoea  alveolaris. 

Entamoeba  undulans^  Aide  Castellani,  1905. 

Under  this  name  a  protozoon  is  described  which  A.  Castellani  found  in  addition 
to  Entamceba  histolytica  and  Trichojtionas  intestinalis  in  the  fasces  of  an  European 
planter  living  in  Ceylon,  who  had  suffered  from  amoebic  enteritis  and  liver  abscess. 
The  shape  of  the  body  was  roundish  or  oval,  25 /t  to  30 /t  in  the  greatest  diameter. 
It  was  without  a  flagellum,  but  with  an  undulating  membrane,  and  capable  of  pro- 
truding a  long  pseudopodium  from  different  parts  of  its  body  at  short  intervals.  The 
nucleus  could  not  always  be  recognized  in  life  ;  it  was,  however,  always  demonstrable 

'  Joiirn.  Amer.  Med.  Assoc,  Ixiv,  p.  553. 


44  THE   ANIMAL   PARASITES   OF   MAN 

by  staining.  One  or  two  contractile  vacuoles  were  present.  The  protoplasm  was 
finely  granular,  showing  no  differentiation  into  ecto-  and  endo-plasm.  According  to 
Braun,  in  spite  of  the  author  declaring  himself  expressly  against  the  flagellate  nature 
of  the  parasite,  such  a  nature  may  be  assumed  to  be  tolerably  certain  in  view  of  the 
description  and  illustration. 

It  is   now   considered    that   Entamcsba    undulans   is    a   portion  of  a  flagellate, 
namely,  T?'ichomonas. 

Entamoeba  kartulisi,  Doflein,  1901. 

Doflein  gave  this  name  to  amoebae,  from  30 /^  to  38 /x  in  diameter, 
which  KartuHs(i893)  found  on  examining  the  pus  of  an  abscess  in  the 
right  lower  jaw  of  an  Arab,  aged  43,  and  in  a  portion  of  bone  that  had  been 

extracted.  The  movements  of  the  amoebae 
(fig.  11)  were  more  active  than  those  of 
^'  dysenteric  amoebae."  Their  coarsely 
granular  cytoplasm  contained  blood  and 
pus  corpuscles,  and  a  nucleus  was  generally 
;  only  recognizable  after  staining.  Vacuoles 
I  ^/'       were  not  seen  with  certainty.     Flexner  re- 

^  ported  upon    a   similar    case,   and   Kartulis 

published  five  additional  cases.  As  in  these 
&rDofl.'; ftotThTpt'oTan  cases  dental  caries  was  present  the  infection 
abscess  in  the  lower  jaw,  show-  is  likely  to  have  proceeded  from  the  oral 
mfnt^'^{AftIr'S-?uliso'"°''^"     cavity   as    a    result   of    the    carious    teeth. 

Craig^  (191 1)  considers  that  this  parasite  is 
probably  identical  with  Entamoeha  histolytica. 

In  the  literature  the  following  species  have  been  reported  as 
occurring  in  the  oral  cavity  of  man  : — 


Amoeba  gingivalis,  Gros,  1849.     [?  identical  with  Eniamoeha  bticcalis.~[ 

Amceba  buccalis,  Sternberg,   1862. 

Amoeba  dentalis,  Grassi,  1879. 

Far  too  little,  however,  is  known  concerning  these  to  regard  them  as  definite 
species,  that  is,  independent  organisms  ;  Grassi  thinks  it  even  possible  there  may 
have  been  a  confusion  in  their  case  with  salivary  corpuscles.  If  they  really  are 
amcebse  they  are  all  of  them  probably  identical  with  E7itamceba  buccalis. 

Genus  Paramoeba,  Schaudinn,   1896. 

Schaudinn  established  the  genus  Paramcsba  for  a  marine  rhizopod  which  multi- 
plied by  division,  became  encysted  at  the  end  of  its  vegetative  life  and  then 
segmented  into  swarm  bodies  with  two  flagella.  These  multiplied  by  longitudinal 
fission,  and  finally  passed  into  the  condition  of  Amoebae.  Whether  the  human  parasite 
described  by  C.  F.  Craig  (1906)  as 

'  "The  Parasitic  Amoebse  of  Man,"  Lippincott,  Philadelphia. 


AMCEBA   UROGENITALIS  45 

Paramoeba  hominis. 

belonged  to  this  genus  was  for  a  time  uncertain.  It  is  now  placed  in  a  new  genus 
Craigia,  Calkins,  1912,  since  it  possesses  only  one  flagellum.' 

In  the  amoebic  stage  it  is  15  ^  to  25  ju  in  diameter  ;  ecto-  and  endo-plasm  during 
rest  are  indistinguishable.  The  body  substance  is  granular,  with  a  spherical,  sharply 
contoured  nucleus  and  an  accessory  nuclear  body.  No  vacuoles  are  present,  but 
occasionally  the  endoplasm  contains  red  blood  corpuscles.  The  pseudopodia  are 
hyaline,  finger-  or  lobe-shaped,  and  are  protruded  either  singly  or  in  twos. 
Multiplication  is  by  binary  fission  and  by  the  formation  of  spherical  cysts  (15/ito 
10  jx  in  diameter)  in  which  occurs  successive  division  of  the  nuclei,  ultimately  forming 
ten  to  twelve  roundish  bodies  each  of  which  soon  develops  a  flagellum.  The  flagellate 
stages  have  similarly  a  spherical  shape  and  attain  a  diameter  of  10  /*  to  15  fi. 
They  also  occasionally  contain  red  blood  corpuscles  and  pass  either  directly  or  after 
longitudinal  division  into  the  amoeboid  phase. 

Craig  found  these  Amoebae  and  the  flagellate  stage  belonging  to  them  in  six 
patients  in  the  military  hospital  at  Manila  (Philippine  Islands),  five  of  whom  were 
suffering  from  simple  diarrhoea  whilst  the  sixth  exhibited  an  amoebic  enteritis  and 
contained  also  Parai7taba  hominis,  with  Entainoeba  histolytica^  Schaudinn.  In  one 
of  the  other  cases.  Trichomonas  intestinalis  was  present. 

B.    Amoebae  from  other  Organs. 

EntamcEba  pulmonalis,  Artault,  1898. 

Arlault^  discovered  a  few  amoeboid  forms  with  nucleus  and 
vacuole  in  the  contents  of  a  lung  cavity.  In  the  fresh  condition  they 
were  distinguishable  from  leucocytes  by  their  remarkable  capacity  of 
light  refraction.  They  were  also  much  slower  than  the  latter  in 
staining  with  methylene  blue  or  fuchsine.  Their  movements  became 
more  lively  in  a  strong  light.  Water  and  other  reagents  killed  them, 
and  then,  even  when  stained,  they  could  not  be  distinguished  from 
leucocytes.  They  have  also  been  seen  by  Brumpt.  R.  Blanchard  found 
amoebae  which  may  belong  here  in  the  lungs  of  sheep.  A.  pulmonalis 
is  perhaps  the  same  as  Entamoeba  buccalis.  Smith  and  Weidman^ 
(1910,  1914)  described  an  entamoeba,  E.  mortinatalmm,  from  the 
lungs  and  other  organs  of  infants  in  America. 

Amoeba  urogenitalis,  Baelz,  1883. 

This  species  was  found  in  masses  in  the  sanguineous  urine  as  well 
as  in  the  vagina  of  a  patient  in  Japan,  aged  23.  Shortly  before  the 
death  of  the  patient,  which  was  caused  by  pulmonary  tuberculosis, 
haematuria  with  severe  tenesmus  of  the  bladder  had  set  in.  The 
amoeba,  which  showed  great  motility,  and  had  a  diameter  of  about 
50  fi  when  quiescent,  exhibited  a  granular  cytoplasm  and  a  vesicular 
nucleus.  Baelz  is  of  opinion  that  these  parasites  were  introduced  into 
the  vulva  with  the  water  used  for  washing  the  parts,  and  thence  had 

'  See  Craig  (1913),  Amer.  Journ.  Trap.  Dis.  and  Prevent.  Med.y  i,  p.  351. 

'  Arch,  de  Parasitologies  i,  p.  275. 

^  A?fier.  fourn.  Trop.  Dis.  and  Prevent.  Med.y  ii,  p.  256. 


46  THE   ANIMAL   PARASITES   OF   MAN 

penetrated  into  the  bladder  and  vagina.     Doflein  places  the  organism 
in  the  genus  Entamoeba,  and  it  is  perhaps  identical  with  E.  histolytica. 

Similar  cases  are  also  reported  (1892-3)  by  other  authors  :  Jiirgens,  Kartulis,  Posner^ 
and  Wijnhoff.  Jiirgens  found  small  mucous  cysts,  filled  with  amoeboid  bodies,  in  the 
bladder  of  an  old  woman  suffering  from  chronic  cystitis ;  they  were  also  found 
in  the  vagina.  The  amoeba  observed  by  Kartulis  in  the  sanguineous  urine  of 
a  woman,  aged  58,  suffering  from  a  tumour  of  the  bladder,  measured  12 /*  to  20  |t, 
and  exhibited  slow  movements  by  protruding  short  pseudopodia.  The  vacuoles  and 
nucleus  became  visible  only  after  staining  with  methylene  blue. 

Posner's  case  related  to  a  man,  aged  37,  who  had  hitherto  been  quite  healthy  and 
had  never  been  out  of  Berlin.  Suddenly,  after  a  rigor,  he  passed  urine  tinged  with 
blood.  This  contained,  besides  red  and  white  blood  corpuscles  and  hyaline  and 
granular  casts,  large  granular  bodies  (about  50 /*  in  length  and  28  /i  in  breath),  which 
slowly  altered  their  shape,  and  contained  red  blood  corpuscles  in  addition  to  other 
foreign  matter.  These  bodies  exhibited  one  or  several  nuclei  and  some  vacuoles. 
From  the  course  of  the  disease,  which  extended  over  a  year,  and  during  which 
similar  attacks  recurred,  Posner  came  to  the  conclusion  that  the  amoebae  which  had 
originally  invaded  the  bladder  had  penetrated  into  the  pelvis  of  the  kidney,  where 
they  probably  had  settled  in  a  cyst,  and  thence  induced  the  repeated  attacks. 

Wijnhoff  observed  four  cases  of  amoeburia  in  Utrecht. 

Amoeba  miurai,  Ijima,  1898. 

Under  this  term  the  author  describes  protoplasmic  bodies  which  Miura,  in 
Tokyo,  found  in  the  serous  fluid  of  a  woman,  aged  26,  who  had  died  from  pleuritis 
and  peritonitis  endotheliomatosa.     Two  days   before  death  these  same  forms  had 

t^-^ —  also  appeared  in   the  haemorrhagic  faeces    of  the 

/  */'""^\  patient.  The  bodies  were  usually  spherical  or 
tr||fafciriBi  ^  .i'-V  \  ellipsoidal,  and  at  one  pole  carried  a  small  pro- 
\m|v',  y:.<,  J/  tuberance  (fig.  12)  beset  with  filamentous  short 
^^jjl^Lm^  "pseudopodia"  (really  a  pseudopodium  covered 
^^^^  with  cilia).     Their    size  varied  between  15  fx  and 

38  /t.      The  cytoplasm    was   finely  granular,    and 

TT.^     ,  y,       L         .       .    T-        1^0     difference     was      observable     in     the     ecto- 

FlG.     12. —  Amceba   imtirai,    Ij.  ,  ,       ,  ,         1  • 

X  500.     a,  fresh  ;     b,   after  treat-       and    endo-plasm,      only    the    villous     appendage 

ment     with     dilute     acetic     acid.       was   clearer.     The  cytoplasm   contained  vacuoles 

(After  Ijima.)  more    or   less    numerous,    none     of     which    was 

contractile.  After  the  addition  of  acetic  acid  one 
to  three  nuclei  could  be  distinguished,  8  ^u  to  15  /t  in  size.  Actual  movements  were 
not  observed.  Taking  everything  into  consideration,  the  independent  nature  of  these 
bodies  is,  to  say  the  least, doubtful,  although  it  cannot  be  denied  that  they  possess  a 
certain  similarity  to  the  marine  Amceba  fluida,  Grilber  or  Greeff,  and  to  a  few  other 
species.   (It  is  likely  that  cells  present  in  serous  exudation  were  mistaken  for  amoebae.) 

Appendix. 

^*  Rhizopods  in  Poliomyelitis  acuta." 

In  three  cases  of  poliomyelitis  acuta  which  were  investigated  by  Ellermann,  the 
spinal  fluid  obtained  by  puncture  of  the  cord  contained  bodies,  from  10 /a  to  15  m  in 
size,  which  had  amoeboid  movements  and  exhibited  variously  shaped  pseudopodia  in 
large  numbers.  After  staining,  a  usually  excentric  nucleus,  about  i"5 /a  in  size,  was 
demonstrated  in  them. 


CHLAMYDOPHRYS    ENCHELYS  47 

Order.     Foraminifera,  d'Orbigny. 

The  order  is  divided  by  Max  Schultze  into  Monothalamia  and  Polythalamia.     Only 
a  few  of  the  former  can  be  considered  here. 

Sub-Order.     Monothalamia.     (Testaceous  Amoebae). 

These  forms  occur  frequently  in  fresh  water,  rarely  in  sea  water. 
They  possess  a  shell  which  is  either  pseudo-chitinous  in  character,  or 
consists  of  foreign  particles,  or  in  a  few  cases  is  composed  of  siliceous 
lamellae.  There  is  usually  an  orifice  for  the  protrusion  of  pseudopodia. 
The  only  representative  of  the  order  of  interest  here  is  : — 

Genus.     Chlamydophrys,  Cienkowski,  1876. 

The  genus  is  based  on  a  form  which  A.  Schneider  carefully  investigated  and 
considered  to  be  the  Dijfflugia  enchelys  of  Ehrenberg.  L.  Cienkowski  redis- 
covered this  same  form  and  created  for  it  the  genus  Chlamydophrys.  We  agree 
with  this  view,  but  not  with  the  renaming  of  the  organism  (so  common  at  the  time). 
If  the  parasite  in  dung,  Chlamydophrys  stercorea  Cienk:  is  identical  with  Difflugia 
enchelys  of  Ehrenberg,  the  old  specific  name  should  be  retained. 

The  genus  is  characterized  by  the  possession  of  a  hyaline,  structure- 
less, slightly  flexible  shell  which  is  ovoid  or  reniform.  At  the  more 
pointed  pole  there  is  an  orifice  situated  terminally  or  somewhat  laterally^ 
serving  for  the  emergence  of  the  filiform  pseudopodia  (fig.  13,  a). 
The  protoplasm  does  not  entirely  fill  the  interior  of  the  shell.  An 
equatorial  zone  bearing  excretory  granules  divides  the  shell  internally 
into  two  almost  equal  portions.  The  anterior  portion  is  rich  in 
vacuoles  and  serves  for  the  reception  of  nutriment  and  for  digestion. 
The  posterior  part  is  vitreous,  and  contains  the  nucleus.  One  to  three 
contractile  vacuoles  are  situated  in  the  equatorial  zone. 

Chlamydophrys  enchelys,  Ehrbg. 
Syn. :  Chlamydophrys  stercorea^  L.  Cienkowski. 

This  species  (fig.  13)  is  found  in  the  faeces  of  various  animals  (cattle, 
rabbits,  mice,  and  lizards),  and  also  in  quite  fresh  human  faeces. 
According  to  Schaudinn,  the  parasite  occurs  so  frequently  in  the 
human  faeces  that  it  must  be  considered  of  wide  distribution.  The 
species  must  traverse  the  intestine  of  man  and  animals  during  one 
stage  of  its  life  cycle,  as  Schaudinn  showed  by  experiments  on  himself 
and  on  mice.  He  infected  himself  with  cysts  (fig.  14)  by  swallowing 
them,  and  evacuated  the  first  Chlamydophrys  as  early  as  the  following 
day.  After  the  evacuation  of  numerous  specimens  on  one  of  the 
following  days  the  infection  ceased. 

The  nucleus  of  a  living  specimen  is  surrounded  by  a  hyaline, 
strongly  retractile  chromidial  mass,  arranged  in  the  form  of  a  ring. 
Chromatin  stains  colour  it  darkly. 

Asexual  inultipHcation  (fig.  13,  /?),  which  takes  place  in  faeces,  follows 


48 


THE   ANIMAL   PARASITES   OF   MAN 


a  similar  course  to  that  of  allied  forms  (e.g.,  Eiiglypha,  Centropyxis).  It 
commences  by  the  cytoplasm  issuing  from  the  orifice  of  the  shell  and 
assuming  the  shape  characteristic  of  the  mother  organism,  but  in 
a  reverse  position.  The  nucleus  then  divides  by  mitosis,  when  the 
daughter  nuclei  move  apart  from  one  another.  The  chromidial  ring 
also  divides  into  two  portions  by  a  process  of  dumb-bell  like  con- 
striction. The  one  daughter  nucleus  remains  in  the  mother  organism, 
the  other  moves  towards  the  daughter  individual,  which  then  separates 
from  the  parent. 


Fig.    13. — Chlamydophrys  ejichelys.       a,    free,    motile   form,    showing   nucleus,    equatorial 
granules,  vacuoles  and  pseudopodia  ;  b,  dividing  organism,      x  760.  (After  Cienkowski.) 


In  this  species  plasmogamic  union  of  two  or  more  individuals  (up  to  twenty)  is 
frequently  observed.  Such  colonies  may  similarly  divide,  and  in  this  way  mon- 
strosities frequently  arise.  When  drying  of  the  fasces,  or  deficiency  of  food  occurs, 
encystment  takes  place  apparently  spontaneously.  The  whole  body,  as  stated  by 
Cienkowski,  issues  from  the  shell,  assumes  a  spherical  shape  (probably  with  discharge 
of  water)  and  becomes  surrounded  with  a  thick  membrane  (fig.  14).  After  the  addition 
of  water  and  the  escape  of  the  encysted  Chlamydophrys^  a  new  shell  must  be  formed. 
Schaudinn,  who  has  not  given  a  more  detailed  description  of  the  process  of  encyst- 
ment in  this  species,  but  refers  to  Cienkowski  and  to  similar  observations  made  on 
Centropyxis^  states  of  the  latter  that  the  encystment  takes  place  within  the  shell. 

The  sexual  nmltiplication  is  accompanied  by  shedding  of  all  the 
foreign  bodies  and  of  the  degenerating  nucleus.  The  protoplasm, 
now  contracting  into  a  sphere,  remains  behind  in  the  shell  with  the 
chromidial  mass.  From  the  latter  several  new  nuclei  arise  (sexual 
nuclei)  often  eight  in  number.  The  cytoplasmic  sphere  then  segre- 
gates into  as  many  spherical  portions  as  there  are  nuclei  present. 
When  they  have  assumed  an  oval  form,  two  flagella  develop  at  one 


LEYDENIA   GEMMIPARA  49 

pole  and  the  flagellispores  swarm  out  of  the  shell/     The  biflagellate 
swarm-spores,  or  gametes,  copulate  in  pairs  and  apparently  the  indi- 
viduals of  the  pairs  of  gametes  arise, from  different  mother  organisms. 
The  zygote  secretes  a  thick  covering  which  soon 
becomes  brown  and  rough.    These  zygote  cysts  >- 

or  resistant  spores  must  now  pass  from  the  in-       /  \ 

testine  of  an   animal  in  order  to  complete  their       (  I 

development.     The  escape  of  the  cyst  contents       1  ^  / 

does  not    always    take  place  in  the  intestine  ;       'v  -  -^=^^^^ 

often  it  does  not  occur  until  after  def?ecaticn.  ^-— ^ 

These  shell-less  individuals  (amoebulae)  soon  ;>;jrj/j  %//il/j7,  ency'Sd''; 
become  invested  with  a  shell.  But  in  the  on  the  left  the  old  capsule, 
alkaline  intestinal  contents,  shell  formation  may      kowskU  ^^      ^^"" 

proceed  even  while  the  organism  is  in  the   in- 
testine, and  multiplication  may  take  place. 

Schaudinn's  further  communication  was  of  special  interest ;  it  was 
to  the  effect  that  Chlamydophrys  was  related  to 


Leydenia  gemmlpara,  Schaudinn,  1896. 

In  the  fluid  removed  by  puncture  from  two  patients  suffering 
from  ascites  in  the  first  medical  clinic  in  Berlin,  cellular  bodies 
with  spontaneous  movement  were  found,  which  Leyden  and 
Schaudinn  regard  as  distinct  organisms.  They  remained  alive 
without  the  use  of  the  warm  stage  for  four  or  live  hours,  the 
external  temperature  being  24°  to  25°  C.  In  a  quiescent  condition 
they  were  of  a  spherical  or  irregular  polygonal  form.  Their  surface  was 
rarely  smooth,  being  beset  with  protuberances  and  excrescences  (fig.  15). 
The  substance  of  the  body  was  thickly  permeated  with  light  refractile 
granules  with  a  yellowish  shimmer.  The  hyaline  ectoplasm  was 
rarely  seen  distinctly.  All  sizes  from  3  /^  to  36  yw,  in  diameter  were 
observed.  The  movements  were  rather  sluggish,  the  ectoplasm  in 
the  meantime  appearing  in  the  form  of  one  or  several  lamellae,  in 
which  also  strings  of  the  granular  endoplasm  occurred,  and  frequently 
protruded  over  the  border  of  the  hyaline  pseudopodia.  The  tendency 
for  the  joining  of  several  individuals  by  means  of  their  pseudopodia 
was  so  marked  that  associations  ensued  similar  to  those  known  in 
free-living  Rhizopoda. 

The  cytoplasm  enclosed  blood  corpuscles  as  well  as  numerous 
vacuoles,  one  of  which  pulsated  slowly  about  every  quarter  of  an 
hour.  A  vesicular  nucleus  the  diameter  of  which  was  about  equal 
to  one-fifth  of  the  body  was  present. 

'  Schaudinn  (1903),  Arb.  a.  d.  Kaiserl.  Gesundh.,  xix,  p.  547. 


50 


THE   ANIMAL   PARASITES   OF   MAN 


Multiplication  took  place  by  means  of  division  andbudding  (lig.  15,  c), 
after  previous  direct  division  of  the  nucleus.  The  buds  were  supposed 
to  divide  repeatedly  soon  after  their  appearance,  thus  giving  rise  to 
minute  forms  of  3  /^. 

There  was  a  suspicion  in  both  cases  that  the  ascites  was  associated 
with  maHgnant  neoplasms  in  the  abdomen,  and  autopsy  confirmed 
this  view  in  one  case. 


c 


X 


Fig.  15. — Leydenia  gemmipara^  Schaud.    a,  in  a  quiescent  condition,  x  looo  ;  ^,  in  the  act  of 
moving,  X  lOOO  ;  f,  from  a  fixed  preparation,  showing  a  bud,  X  1500. 


The  parasite,  which  has  seldom  been  observed,  has  been  variously 
interpreted ;  for  example,  it  has  been  regarded  merely  as  altered  tissue 
cells.  It  is  now  known,  from  Schaudinn's  researches,  that  Leydetiia 
geminipara  is  connected  with  abnormal  conditions  of  Chlainydophrys, 
occasionally  occurring  as  a  commensal  in  the  ascitic  fluid.  The  form 
is  produced  when  pathological  conditions  of  the  large  intestine  create 
an  alkaline  reaction  of  its  whole  contents.  The  formation  of  shells 
then  often  ceases,  and  these  naked  Chlaniydophrys  are  enabled  to 
multiply  atypically  by  division  and  gemmation.  Such  stages,  which 
are  no  longer  capable  of  a  normal  development,  are  the  Leydenia,  as 
Schaudinn  has  demonstrated. 


Class  II.     MASTIGOPHORA,  Diesing. 
Sub-Class.     FLAGELLATA,  Cohn  emend.  Biitschli. 

During  the  motile  part  of  their  life  the  Flagellata  possess  one  or  more  fiagella 
which  serve  for  locomotion,  and  in  many  cases  also  for  the  capture  of  food.  A  few 
groups  {Euglenoidi?2CEy  Choanofla^ellata)  have  only  one  fiagellum,  others  two  or 
several  of  about  equal  length  {Isomasiigoda)^  or  of  various  lengths  {Monadina, 
Heteromastigoda^  Dinofiagellata).  The  long  llagellum  is  the  principal  one  ;  the 
smaller  ones  on  the  same  organism  are  accessory  fiagella.     The  flagella  directed 


MASTIGOPHORA  5 1 

backwards,  which  occur  in  the  Heteromastigoda  and  are  used  for  clinging,  are 
termed  trailing  flagella  or  tractella.  At  the  base  of  the  flagellum,  which  is  almost 
always  at  the  anterior  end,  a  Choanoflagellate  possesses  a  cytoplasmic  funnel-shaped 
neck  or  collar.     In  the  parasitic  forms  an  undulating  membrane  is  often  present. 

The  body  of  the  Flagellata  is  usually  small,  generally  elongate  and  of  un- 
changeable form.  It  is  frequently  covered  by  a  distinct  cuticle,  and,  in  certain  groups, 
by  a  hard  envelope,  or  it  may  be  more  or  less  loosely  enveloped  by  a  gelatinous  or 
membranous  covering.  An  ectoplasmic  layer  is  thin  and  not  always  obvious.  The 
granular  cytoplasm  contains  a  varying  number  of  vacuoles,  one  of  which  may  be 
contractile,  and  is  generally  situated  near  the  area  from  which  the  flagella  arise, 
that  is,  at  the  anterior  extremity.  The  cytoplasm,  moreover,  contains  the  nucleus, 
which  is  nearly  always  single  ;  and  in  many  species  there  are  also  yellow,  brown, 
or  green  chromatophores  of  various  shapes,  such  as  occur  in  plants.  Some 
species  feed  after  the  manner  of  green  plants  (holophytic),  or  of  plants  devoid  of 
chlorophyll  (saprophytic)  ;  others,  again,  ingest  solid  food,  and  for  this  purpose 
usually  possess  a  cytostome  ;  the  latter,  however,  in  a  few  forms  is  not  used  for  its 
original  function,  but  is  connected  with  the  contractile  vacuole.  Many  parasitic 
forms  feed  by  endosmosis.  A  few  species  possess  eye-spots  with  or  without  light- 
refracting  bodies. 

Variation  in  the  form  of  the  nuclear  apparatus  occurs.  One  nucleus  only,  which 
may  be  compact  or  vesicular,  is  known  in  many  species.  This  nucleus  is  situated 
either  centrally  or  sometimes  near  the  flagellar  end  of  the  body,  but  its  position  is 
subject  to  variation.  The  flagella  may  arise  near  the  nucleus.  Other  structures, 
such  as  an  axial  filament  and  a  rhizoplast,  may  be  present.  Some  flagellates  are 
binucleate,  the  two  nuclei — which  often  differ  in  size  and  shape — being  separated 
from  each  other.  One  of  these  nuclei  is  the  principal,  vegetative  or  trophic  nucleus  ; 
the  other  is  an  accessory  nucleus,  frequently  termed  the  blepharoplast,  flagellar 
or  kinetic  nucleus.  One  or  more  small  basal  granules  are  often  present  at  or  very 
near  the  origin  of  the  flagella. 

Multiplication  is  by  fission,  usually  longitudinal,  which  may  occur  in  either  the 
free  or  encysted  forms.  Division  is  initiated  by  that  of  the  nucleus  or  nuclei 
(especially  the  kinetic  nucleus).  The  basal  granule  divides  also.  Collars  and 
chromatophores,  if  present,  likewise  separate  into  two.  Variation  in  the  method  of 
doubling  the  original  number  of  flagella  occurs.  In  most  organisms,  especially 
uniflagellate  forms,  the  flagellum  splits  lengthwise,  after  division  of  the  basal  granule, 
blepharoplast  and  nucleus.  The  daughter  flagella  may  be  of  the  same  or  different 
lengths  and  thicknesses.  Other  flagellates  at  division  are  said  to  produce  new 
flagella  in  the  neighbourhood  of  the  original  ones.  The  daughter  organisms  in  such 
cases  are  provided  with  one  or  more  parental  flagella  in  addition  to  newly  formed 
ones.  It  has  been  stated  that  in  certain  cases  the  parent  flagellate  retains  all  its 
flagella,  while  new  ones  arise  ab  initio  in  the  cytoplasm  of  the  daughter  forms. 

Multiplication  by  longitudinal  fission  may  be  interrupted  sooner  or  later  by  the 
production  of  gametes,  which  form  zygotes,  from  which  new  generations  of  indi- 
viduals arise.  In  many  flagellates  gamete  formation  and  sporogony  are  unknown, 
and  asexual  reproduction  by  fission  alone  prevails. 

Incomplete  division  results  in  the  formation  of  colonies  of  individuals.  These 
colonies  must  not  be  confused  with  the  aggregation  rosettes  of  flagellates  found 
among  the  parasitic  Mastigophora.  The  individuals  of  aggregation  rosettes  are 
capable  of  immediate  separation  from  the  rosette  at  will. 

A  number  of  parasitic  Flagellata  produce  non-flagellate  stages  which  are  very 
resistant  to  external  conditions,  the  assumption  of  which  forms  serves  to  protect  the 
organisms  during  their  transference  from  one  host  to  another.  Such  non-fla^ellate 
forms  possess  one  or  more  nuclei,  are  usually  of  an  oval  or  rounded  contour,  and 


52  THE   ANIMAL   PARASITES   OF   MAN 

are  capable  of  developing  into  the  full  flagellate  on  the  return  of  more  favourable 
conditions.  These  forms  are  often  known  as  the  post-flagellate  stage  of  the 
organism.  When  ingested  by  a  new  host,  the  post-flagellate  coat  becomes  more 
flexible,  and  the  phase  of  the  organism  which  now  recommences  growth  is  known  as 
the  pre-flagellate  stage  ;  it  gradually  develops  into  the  typical  flagellate  organism. 

Many  Flagellata  live  free  in  fresh  and  salt  water.  They  prefer  stagnant  water, 
rich  in  organic  products  of  decomposition,  such  as  puddles,  swamps  and  pools. 
Those  forms  developing  shells  and  colonies  are,  as  a  rule,  adherent.  A  number  of 
species  are  parasitic  in  man  and  animals,  living  mostly  within  the  intestine  or  in  the 
blood. 

It  is  usual  to  classify  the  Flagellata  in  four  orders  :  Euflagellata,  Dinoflas^ellata^ 
Choanoflagellata^  and  Cysto flagellata^  of  which  only  the  Euflagellata  are  of  mterest 
to  us.  This  is  a  group  comprising  numerous  species,  for  the  further  classification 
of  which  the  number  and  position  of  the  flagella  are  utilised. 

The  Euflagellata  observed  in  man  belong  to  the  Protomonadina  as  well  as  to  the 
Polymastigina.  The  former  possess  either  only  one  or  two  similar  flagella,  or  one 
principal  and  one  or  two  accessory  flagella.  The  Polymastigina  possess  at  least 
three  flagella  of  equal  size,  or  four  to  eight  of  unequal  size,  inserted  at  different 
points.     An  undulating  membrane  may  be  present  in  members  of  both  groups. 

It  must  also  be  pointed  out  that  unicellular  organisms  with  one  or  several 
flagella  are  not  always  classified  with  flagellates,  for  such  forms  occur  in  Rhizopods 
as  well  as  temporarily  in  the  lower  plants.  In  addition,  the  examination  of  the 
flagellates,  especially  the  parasitic  species,  is  very  difficult  on  account  of  their 
diminutive  size  and  great  activity  ;  thus  it  happens  that  certain  forms  cannot  with 
certainty  be  included  in  the  group  because  their  description  is  insufficient. 

Order.     Polymastigina,  Blochmann. 

The  Polymastigina  contains  flagellates  with  three  to  eight  flagella. 
Some  of  the  Flagellata  parasitic  in  man  belong  to  the  Polymastigina, 
and  to  two  or  three  genera  that  are  easily  distinguishable. 

Genus.     Trichomonas,  Donne,  1837. 

The  body  is  generally  pyriform,  the  anterior  part  usually  rounded,  the  posterior 
part  pointed.  There  are  at  the  anterior  extremity  three  (?  four)  equally  long  flagella 
that  are  sometimes  matted  together.  A  blepharoplast  (kinetic  nucleus)  and  basal 
granule  are  present,  together  with  a  supporting  structure  known  as  an  axial 
filament  or  axostyle.  In  addition  there  is  an  undulating  membrane,  bordered  by  a 
trailing  flagellum,  that  commences  at  the  anterior  extremity  and  proceeds  obliquely 
backwards.  The  nucleus,  which  is  vesicular,  is  situated  near  the  anterior  extremity, 
and  behind  it  are  one  or  more  vacuoles,  none  of  which  seems  to  be  contractile. 
These  flagellates  are  parasitic  in  vertebrate  animals,  and  live  chiefly  in  the  intestine. 

Trichomonas  vaginalis,  Donne. 

The  form  of  the  body  is  very  variable,  and  is  elongate,  fusiform 
or  pear-shaped,  also  amoeboid.  The  length  varies  between  15/1,  and 
25  /i,  and  the  breadth  between  7  ytt  and  12  /i.  The  posterior  extremily 
is  drawn  out  to  a  point  and  is  about  half  the  length  of  the  remainder 


TRICHOMONAS   VAGINALIS 


53 


of  the  body.  The  cuticle  is  very  thin  and  the  body  substance  finely 
granular.  At  the  anterior  extremity  there  are  three — some  say  four  ^ — 
flagella  of  equal  length  which  are  frequently  united  together,  at  least 
at  the  base,  and  are  easily  detached. 

There  is  an  undulating  membrane  (fig.  i6) 
which  runs  spirally  across  the  body,  arising 
from  the  place  of  insertion  of  the  flagella,  and 
terminating  at  the  base  of  the  caudal  process. 
A  cytostome  seldom  is  recognizable  in  fresh 
specimens,  but  is  apparently  present.  The 
nucleus  is  vesicular,  elliptical  and  situated  near 
the  anterior  extremity.^ 

Multiplication  takes  place  by  division 
(Marchand).  Encysted  forms  are  almost  un- 
known. 

Trichomonas  vaginalis  lives  in  the  vaginal  mucus  of 
vvomen  of  various  ages,  not  in  normal  mucus,  but  in 
mucus  of  acid  reaction.  It  is  found  in  menstruating 
females  as  well  as  in  females  who  have  passed  the 
menopause.  It  occurs  in  pregnant  and  non-pregnant 
women,  even  in  very  young  girls,  provided  always  that 
they  have  a  vaginal  catarrh  with  acid  reaction  of  the 
secretion.  Should  the  acid  reaction  change,  as,  for 
instance,  during  menstruation,  the  parasites  disappear, 
as  they  do  likewise  on  injection  of  any  alkaline  fluid  into 
the  vagina.  A  low  temperature  (below  +  15°  C.)  is  also 
fatal  to  the  parasites.  These  flagellates  can  pass  from 
the  vagina  through  the  urethra  into  the  bladder,  and 
produce  severe  catarrh,  and  are  not  easily  removed. 

r.  vaginalis  appeared  to  be  a  parasite  specific  to  the  female  organs 
and  not  transmissible  to  man.  However,  several  observations  have 
since  been  made  that  confirm  the  occurrence  of  this  species  in  the 
urethra  of  the  male.  The  infection  apparently  takes  place  through 
coitus  when  changes  are  present  in  the  urethral  mucous  membrane. 
At  any  rate,  three  cases  observed  point  to  this  circumstance. 

Attempts  at  experimental  transmission  to  rabbits,  guinea-pigs  and 
dogs  failed  (Blochmann,  Dock).  So  far,  the  manner  in  which  women 
become  infected  is  unknown. 


Fig.  16. —  Trichomonas 
vaginalis,  Donne.  X  2,000 
approx.  (After  Kunstler.) 
Four  flagella  are  repre- 
sented, but  usually  only 
three  are  present. 


1  To  explain  this  discrepancy  it  is  stated  that  the  border  of  the  undulating  membrane  can  be 
detached  in  the  form  of  an  independent  flagellum.  But  Parisi  (1910)  places  such  quaciriflagellate 
forms  in  the  sub-genus  Teiratrichomonas ,  Arch.  f.  Protistenk.,  xix,  p.  232. 

■^  According  to  Marchand,  the  nucleus  is  connected  with  a  line,  which  becomes  visible  on 
addition  of  acetic  acid,  terminates  at  the  posterior  extremity,  and  does  not  correspond  to  the 
line  of  insertion  of  the  undulating  membrane.  This  formation  probably  is  the  same  as  the 
axostylein  Trichomonas  bairacho?'ttm,  Perty.  Blochmann  (1884)  also  mentions  two  longitudinal 
rows  of  granules,  which  commence  at  the  same  place  as  the  nucleus  and  converge  posteriorly. 


54 


THE   ANIMAL   PARASITES   OF   MAN 


Trichomonas  intestinalis,  R.  Leuckart,  1879=  Trichomonas 
hominis,  Davaine,  1854. 

Some  authors  believe  that  a  second  trichomonad  inhabiting  man, 
Trichomonas  intestinalis,  R.  Lkt.,  is  identical  with  Trichonionas  vaginalis, 
Donne.  Leuckart's  species  was  based  on  the  discoveries  of  Marchand 
(1875)  and  Zunker  (1878),  who  stated  that  according  to  all  appearances, 
and  in  their  opinion,  it  was  the  same  as  Cercomonas  intestinalis,  Lambl, 

1875  {nee  1859),  which  they  found  in 
the  fasces  of  patients  suffering  from 
intestinal  disorders.  The  organism 
is  described  by  them  as  being  pear- 
shaped  and  10  yLt  to  15  yLt  in  length 
and  3  /x  to  4'  yLt  in  breadth.  The 
posterior  extremity  terminated  in  a 
point  (fig.  17). 


Fig.    17. 


Trichomonas  intestinalis, 
(After  Grassi.) 


A  row  of  twelve  or  more  cilia  was  said  to 
commence  at  the  anterior  end  and  extend 
Lkt.  over  the  body.  Leuckart  stated  that  this 
parasite,  placed  by  the  two  authors  in  the 
genus  Cerconw?ias\  w^s  a  Trichomonas,  and 
that  they  mistook  the  undulating  membrane  for  cilia,  and  overlooked  the  flagella. 
Notwithstanding  its  striking  similarity  with  T.  vaginalis,  it  was  said  to  be  distin- 
guishable from  that  species  by  ditferences  in  the  undulating  membrane.  Lambl's 
C.  intestinalis^  (of  1875)  which  corresponds  with  C.  hojnifiis,  Davaine^  (1854),  is 
regarded  by  Leuckart  as  a  true  Cercomonad  (characterized  by  a  flagellum  and  the 
absence  of  an  undulating  membrane,  see  p.  61),  and  is  thus  generically  distinct 
from  Trichomonas. 

The  correctness  of  Leuckart's  judgment  in  regard  to  Marchand-Zunker's  flagellate 
was  demonstrated  by  Grassi's  researches,  accounts  of  which  were  published  soon 
after.  In  about  100  cases  of  bowel  complaints  in  North  Italy  and  Sicily,  Grassi 
found  Flagellata  in  the  stools,  which  he  first  named  Monocercomo7tas  and  Ci?ncr?io- 
monas,  but  later  termed  Trichomonas.  However,  in  opposition  to  Leuckart,  Grassi 
has  also  classified  Uavaine's  C.  homi7iis  {=  C.  intestinalis,  Lambl,  1875)  ^s 
Trichomonas,  and  most  authors  have  followed  his  example.  Hence  arose  the  use 
of  the  name  Trichomonas  hominis.  It  was  through  Janowski  (1896)  that  the 
former  view  was  agam  taken  up.  After  a  review  of  the  literature,  the  occur- 
rence of  Cercomonads  in  the  intestine  of  human  beings  in  addition  to  Tricho- 
monads  was  considered  by  the  author  to  have  been  proved,  and  he  added  a 
description  of  the  Trichomonads.     According  to  this,  all  morphological  distinction 


1  Under  the  term  Cercomonas  intestinalis,  Lambl  in  different  years  has  described  two 
entirely  distinct  Flagellata,  namely,  in  1859  ("Mikr.  Unters.  d.  Darm-  Excrete,"  Pnag. 
Vierteljahrsschr.  f.  prakt.  Hlkde.,  Ixi,  p.  51  ;  and  Lambl,  A.  d.  Franz-Josephs- Kinder spitale 
in  Prag,  Prag,  i860,  i,  p.  360),  a  form  that  at  the  present  day  is  termed  Lainblia  intes- 
tinalis;  and  in  1875  (^^^  the  Russian  Medical  Report,  No.  33),  a  species  identical  with 
Cercomonas  hominis,  Dav. 

^Davaine,  C. ,  *' Sur  les  anim.  infus.  trouv.  dans  les  selles  d.  malad.  atteints  du  cholera 
et  d'autr.  malad.,"  C.  R.  Soc.  Biol,,  1854,  ii,  p.  129. 


TRICHOMONAS    INTESTINALIS 


55 


between  T.  vaginalis^  Donne,  and  T.  intestmalis^  LeuckaU,  disappeared.  On  the 
other  hand,  it  is  worthy  of  note  that  the  smaller  size,  the  more  pear-shaped  form,  and 
the  longer  flagella  differentiate  T.  intestinalis  (=  T.  hommis)  from  T.  vaginalis} 

The  easily  deformed  pear-shaped  body  has  three  free  flagella 
anteriorly,  and  an  undulating  membrane  with  its  flagellar  bordc" 
terminating  in  a  short  free  flagellum  posteriorly  (figs. 
17,  18).  The  undulating  membrane  may  coil  itself 
spirally  round  the  body.  A  supporting  rod  or  axo- 
style  projects  as  a  posterior  spine.  It  appears  to 
begin  near  the  nucleus  and  blepharoplast,  which  are 
situated  near  the  more  rounded,  anterior  end  of  the 
body.  There  may  be  a  chromatoid  basal  supporting 
line  along  the  body  for  the  undulating  membrane. 
Rows  of  chromatoid  granules  are  sometimes  situated 
along  one  side  of  the  axostyle.  A  cytostome  may 
sometimes  be  seen.  In  mice,  Wenyon  (1907)  found 
these  parasites  to  vary  in  length  from  3  //,  to  20  \i. 
They  occur  in  the  caecum  and  intestine  of  mice, 
where  their  internal  structure  seems  more  obvious 
than  in  man.  The  flagellates  divide  by  longitudinal 
fission. 

T.  intestinalis,  R.  Leuckart,  appears  to  be  capable 
of  settling  in  all  parts  of  the  human  intestine  in  which 
the  contents  have  an  alkaline  reaction.  Trichomonads 
have  been  cited  as  occurring  in  the  oral  cavity  by  depressior°^anter- 
Steinberg,  Zunker,  Rappin  and  Prowazek ;  in  the  ioriy,  undulating 
oesophagus  by  Cohnheim,  and  in  the  storpach  by  ^^  a^xos^y^l"! 
Strube,  Cohnheim,  Zabel,  Hensen  and  Rosenfeld.  x  2,500.  Original. 
The  normal  situation  seems  to  be  the  small  intestine. 
The  parasites  then  appear  in  the  dejecta,  especially  in  various  intestine, 
diseases  the  course  of  which  is  connected  with  an  increased  peristalsis. 
They  are  also  found  in  healthy  persons,  from  whom  they  are  obtained 
after  the  administration  of  laxatives.  They  have  been  regarded  by  some 
workers  as  commensals,  which,  however,  have  the  power  of  accelerating 

1  For  the  present  the  following  should  be  regarded  as  synonymous  :  Protoryxomyces 
coprinarius^  Cunningham  {Quart.  Journ.  Micr.  ScL  [2]  1880,  xxi,  p.  234),  {Zeitschr.  f.  Biol., 
1882,  viii,  p.  251).  Monocercomorias  hominis,  Grassi,  1882.  Cimcenomonas  hominis^Qx2&%^_ 
1882.  Trichomonas  hominis,  Grassi,  1888.  Cercomonas  coli  hominis^  May  {DetUschei 
Archiv.  f.  klin.  med.,  1 89 1,  xlix,  p.  51).  Monocercomonas  kominis,  Epstein  [Frag.  med. 
Wochenschr.  1893,  ^^s.  38-40).  Trichomonas  confusa,  Stiles  {Zool.  Anz.,  1902,  xxv,  p.  689). 
Trichomonas  elongata.  Trichomonas  elliptica,  Cohnheim  {Deutsche  med.  Wochenschr.,  1903 
xxix,  Nos.  12-14).  Trichomonas  elotigata.  Trichomonas  caudata,  Trichomonas  Jlagellata, 
Steinberg  (A7^ze/(?r  Zeitschr.  f.  neiiere  A/edicin,  1862).  Trichomonas pulmonalis,  A.  Schmidt, 
{Miinch.  med.  Wochenschr.,  1895,  No.  51),  and  St.  Artault  {Arch,  de  parasit.  1898,  i, 
p.   279). 


Fig.  18.— 7>/- 
chomonas  intestin- 
alis from  man, 
showing      anterior 


50  THE   ANIMAL   PARASITES   OF   MAN 

the  onset  of  intestinal  complaints,  or  at  least  of  adding  to  them.  They 
have  been  found  in  cases  of  carcinoma  of  the  stomach,  and  in  other 
diseases  of  that  organ  in  which  the  acid  reaction  ceased. 

Naturally,  whether  all  the  reports  relate  to  the  same  species  of  Trichomonas 
must  remain  undecided.  Certain  authors  (Steinberg,  Cohnheim,  van  Emden) 
accept  several  species.  Provvazek  speaks  of  a  variety  of  T.  intestinalis  inhabiting 
the  oral  cavity.  This  was  distinguished  by  a  posterior  process  exceeding  the 
length  of  the  body  fourfold,  and  by  a  somewhat  unusual  course  of  the  undulating 
membrane.  The  food  of  this  form,  which  was  found  in  the  whitish  deposit  present, 
especially  in  the  cavities  of  carious  teeth,  consisted  almost  exclusively  of  micrococci. 
Schmidt  and  St.  Artault  named  the  Trichomonads  found  in  pathological  products 
{e.g.^  gangrene,  putrid  bronchitis,  phthisis)  of  the  lungs  of  man,  as  Trichomonas 
pulmonalis.  Trichomonads  have  also  been  found  by  VVieting  in  lobular  pneumonia 
in  the  lungs  of  pigs. 

It  is  still  uncertain  in  what  way  the  infection  takes  place.  Experiments  in  the 
transmission  of  free  trichomonads  to  mammals  [per  os)^  in  which  the  same  or  allied 
species  occur  (guinea-pigs,  rats,  apes),  have  been  without  result.  Probably  encyst- 
ment  is  necessary.  Such  conditions  are  mentioned  by  May,  KUnstler,  Roos, 
Schurmayer,  van  Emden,  Prowazek,  Galli-Valerio  and  Schaudinn.  According  to 
Prowazek,  intestinal  trichomonads  of  rats  become  encysted  for  conjugation.  In  the 
cyst  an  accumulation  of  reserve  food  material  occurs,  causing  distension.  The  nuclei 
of  the  conjugants  each  give  off  a  reduction  body  and,  after  fusion,  produce  the 
nuclei  for  the  daughter  individuals.  According  to  Schaudinn  the  intestinal  tricho- 
monads lose  their  flagella  before  conjugation,  become  amoeboid  and  encyst  in  twos, 
the  formation  of  a  large  agglomeration  of  reserve  substance  accompanying  this. 
Galli-Valerio  found  double-contoured  cysts  in  the  faeces  of  trichomonad-infected 
guinea-pigs,  after  the  faeces  had  been  kept  for  a  month  in  a  damp  chamber.  When 
exposed  to  heat  small  flagellates  escaped  from  them.  Administration  of  such  material 
containing  cysts  resulted  in  severe  infection  with  trichomonads,  and  death  of  the 
experimental  guinea-pigs  followed.  The  cyst  wall  is  clearly  a  protection  against  the 
deleterious  acid  reaction  of  the  stomach  contents.  Alexeieff  (1911)  and  Brumpt  (1912) 
think  that  the  trichomonad  cysts  of  man  are  really  fungi,  while  other  workers  also 
doubt  encystment  among  trichomonads.  Wenyon  (1907)  states  that  T.  intestinalis  in 
mice  produces  spherical  contracted  forms  which  escape  from  the  body  in  the  faeces. 

Air,  water,  and  under  certain  circumstances  even  food  may  be  regarded  as 
vectors  for  the  trichomonads.  The  occurrence  of  the  organisms  in  the  oral  cavity, 
and  still  more  so  in  the  lungs,  is  in  favour  of  the  air  being  the  transmitting  agent. 
An  observation  made  by  Epstein  supports  the  idea  of  water  transmission.  Multipli- 
cation of  the  trichomonads,  once  they  have  gained  access  to  the  body,  is  effected  by 
longitudinal  division  commencing  at  the  anterior  end  (Kiinstler).  "  Cercomonads  " 
with  several  flagella  and  an  undulating  membrane,  as  well  as  trichomonads,  have 
been  observed  by  Ross  in  some  cases  of  cutaneous  ulcers. 

Mello-Leitao  (1913)^  has  described  flagellate  dysentery  in  children 
in  Rio  de  Janeiro.  He  states  that  it  is  due  to  T.  intestinalis 
and  Lamhlia  intestinalis  either  separately  or  together.  Flagellate 
dysentery,  he  thinks,  is  benign  and  is  the  most  frequent  form  of 
dysentery  in  infants.  The  flagellates  are  pathogenic  to  infants  under 
three  years  of  age.  Escomel  (1913)^  found  152  cases  of  dysentery  in 
Peru  due  solely  to  Trichomonas.    Such  cases  are  probably  widespread 

'  BriL  Jow-n.  Childretis  Diseases,  x,  p.  60.  ^  jguii.  Soc.  Path.  Exot.,  vi,  p.  120. 


TETRAMITUS   MESNILI  57 

Genus.     Tetramitus,  Perty,  1852. 

Tetramltus  mesnili,  Wenyon,  1910. 
Syn. :   Macrostoma  mesnili^  Chilomastix  mesnili^  Fanapapea  intestinalis. 

The  genus  Tetramitus  differs  from  Trichomonas  in  possessing  an 
undulating  membrane  inserted  in  a  deep  groove  or  cytostome.  There 
are  three  anterior  flagella.  The  pear-shaped  organism  measures  14  yu 
by  7  yLt,  but  smaller  examples  occur.  T.  mesnili  occurs  in  the  human 
intestine,  having  been  described  by  Wenyon^  (1910)  from  a  man  from 
the  Bahamas  in  the  Seamen's  Hospital,  London.  Its  occurrence 
is  widespread.  Alexeieff  considers  that  Macrostoma  and  Tetramitus 
are  synonymous.  The  parasite  is  the  same  as  Fanapapea  intestinalis, 
Prowazek,  1911,  from  Samoa.  Brumpt  (191 2)  found  T.  mesnili  to  be 
the  causal  agent  of  colitis  in  a  Frenchwoman.  Nattan-Larrier  (1912) 
considers  it  of  little  pathological  importance. 

GabeP  (1914)  described  an  interesting  case  of  seasonal  diarrhoea 
acquired  in  Tunis,  in  which  a  new  Tetramitid  was  the  causal  agent. 
The  organism  was  pear-shaped,  without  an  undulating  membrane, 
and  measured  6*5  //,  to  8  //,  by  5  //.  to  6  /x.  The  cytostome  was  large, 
and  there  was  no  skeletal  support.  Encystment  occurred.  Gabel 
named  the  organism  Difdmns  tiinensis  and  considered  that  it  was 
pathogenic. 

Genus.     Lamblla,  R.  Blanchard,  1888. 

Syn.,  Dimorphus,  Grass),  1879,  ^^<^  Haller,  1878;  Me^astoma,  Grassi,  1881,  nee 
de  Blainville. 

The  body  is  pear-shaped,  with  a  hollow  on  the  under  surface  anteriorly.  It  has 
four  pairs  of  tlagella  directed  backwards,  of  which  three  pairs  lie  on  the  borders  of 
the  hollow  disc,  and  the  fourth  arises  from  the  pointed  posterior  extremity. 

Lamblia  intestinalis,  Lambl,  1859. 

Syn.:  Cercomonas  intestinalis^  Lambl,  1859  {nee  1875);  Hexamitus  duodenalis, 
Davaine,  1875;  Dimorphus  inwris,  Grassi,  1879;  Megastoma  enterieum^  Grassi, 
1881  ;   Megastoma  intestinale,  R.  Blanch.,  1886;  Lamblia  duodenalis^  Stiles,  1902. 

The  organism  is  pear-shaped  and  bilaterally  symmetrical.  It  is 
from  10//.  to  21 /z  long  and  5  yw,  to  12  yu,  broad  and  possesses  a  thin 
cuticle.  Anteriorly  an  oblique  depression  is  present,  which  functions 
as  a  sucking  disc  (fig.  19,  s).  Its  edges  are  raised  above  the  general 
surface  and  are  contractile.  It  corresponds  to  a  peristome  and  acts  as 
an  adhesive  organ  (fig.  20,  6,  c).  No  true  cytostome  is  present.  A 
double  longitudinal  ridge,  representing  axostyles,  extends  from  the 
sucking  disc  to  the  tapering  posterior  extremity,  which  is  prolonged 
as  two  flagella  from  9  /x  to  14  /^  long. 

Lamblia  intestinalis  possesses  eight  flagella  (fig.  19).     The  first  pair 

1  Parasitology,  iii,  p.  210.  '^  Arch.  f.  Protistenk.,  xxxiv,  p.  I. 


58 


THE   ANIMAL   PARASITES   OF   MAN 


of  flagella,  which  cross  one  another,  arise  in  a  groove  formed  by  the 
anterior  edge  of  the  sucking  disc.  Two  pairs  of  flagella  (lateral  and 
median)  are  inserted  on  the  posterior  edge  of  the  disc,  while  the 
posterior  flagella  occur  at  the  tapering  posterior  extremity  of  the  body. 
Basal  granules  are  found  at  the  bases  of  the  flagella.  The  median 
flagella  are  most  active  in  movement,  the  anterior  and  lateral  flagella 
being  less  motile,  as  they  are  partially  united  to  the  body  for  part  of 
their  length. 

The  nuclear  apparatus  is  situated  in  the  thin,  anterior,  hollowed 
part  of  the  body.  It  is  at  first  dumb-bell  shaped,  the  "  handle  "  of  the 
dumb-bell  being  formed  by  a  very  slight  connecting  strand,  which 
eventually  separates,  so  that  the  flagellate  becomes  binucleate,  and  thus 
completes  the  general  bisymmetry  of  the  organism. 

There  is  a  karyosome  in  each  nucleus.  Other  bodies  of  unknown 
function,  and  possibly  composed  of  chromatin,  occur  on  or  near 
the  axostyles. 


.  Fig.  19.  —Lamblia  intestinalis.  A,  ventral  view  ;  B,  side  view ;  N,  one  of  the  two  nuclei ; 
ax.^  axostyles;  fl'^,Jl'^,Jl^,fl^,  the  four  pairs  of  flagella;  s,  sucker-like  depressed  area  on  the 
ventral  surface;  x,  bodies  of  unknown  function.     (After  Wenyon.) 


Division  has  not  been  observed  in  the  flagellate  stages  of  the 
Lamblia,  but  it  occurs  within  the  cysts.  The  resistant  cysts  (fig.  20,  e) 
are  oval  and  are  surrounded  by  a  fairly  thick,  hyaline  cyst  wall. 
They  measure  10  //,  to  15  /x  by  7  //<  to  9  /x,  and  may  be  tetranucleate. 
According  to  Schaudinn,  the  cysts  arise  from  the  conjugation  of  two 
individuals,  and  nuclear  rearrangement  occurs. 


LAMBLIA   INTESTINALIS 


59 


L.  intestinalis  occurs  in  its  flagellate  stage  in  the  duodenum  and 
jejunum,  and  rarely  as  such  in  the  other  parts  of  the  intestine. 
Normally  it  is  found  in  the  large  intestine  as  cysts,  which  are  voided 
with  the  faeces.  The  hosts  of  Lamblia  include  Mtis  muscuhis,  M. 
rattuSy  M.  decumanus,  M.  silvestris,  Arvicola  arvensis  and  A,  amphihius 
the  dog  and  cat,  rabbit,  sheep  and  man.  Cysts  voided  with  the  faeces 
of  infected  animals  reach  plants  or  drinking  water,  and  thence  are 
transferred  to  man. 

The  flagellate  in  these  different  hosts  exhibits  some  variation  in 
size  and  in  the  problematic  chromatic  bodies.  Bensen  has  suggested 
the  species  L.  intestinalis  from  man,  L.  ninris  from  the  mouse  and 
L.  cunicnli  from  the  rabbit.  It  is  not  certain  whether  these  different 
species  are  necessary,  as  the  variation  may  be  due  to  differences  of 
environment. 


Fig.  20. — Lamblia  intestinalis.     a,  from  the  surface  ;  b,  from  the  side  ;  c,  on  intestinal 
epithelium  cells  ;  d,  dead  and  <?,  encysted.     (After  Grassi  and  Schewiakoff.) 


Like  Trichomonas,  Lamblia  can  multiply  under  inflammatory 
conditions  of  the  alimentary  tract.  Thus  they  are  found  in  cases 
of  diarrhoea,  carcinoma  of  the  stomach,  etc.  The  parasites  attach 
themselves  by  their  sucking  discs  to  the  epithelial  cells  of  the  gut 
{fig.  20,  c),  and  though  their  numbers  may  be  very  great,  their  direct 
pathological  significance  is  not  fully  known.  Their  occurrence  in 
cases  of  diarrhoea  has  been  explained  as  being  due  to  the  increased 
peristalsis,  which  has  detached  the  parasites  from  the  epithelium. 
Free  flagellate  forms  perish  in  stools  if  kept,  more  especially  if  the 
temperature  falls  below  0°  C.  or  rises  above  40°  C.  Lamblia  has 
often  been  found  in  dysenteric  diseases,  especially  in  the  East,  and 
is  said  to  be  the  causal  agent  of  certain  diarrhoeas  in  India.  Math  is 
{1914)^  found  Lamblia  in  cases  of  diarrhoea  with  dysenteriform  stools 
in  Tonkin.     He  also  discovered  healthy  carriers  of  Lamblia  cysts. 


Bull.  Soc.  Med.  Chirurg.  Indo-Chine,  v,  p.  55. 


6o  THE   ANIMAL   PARASITES   OF   MAN 

The  parasite  under  discussion  was  first  observed  by  Lambl  (1859)  in  the  mucous 
evacuations  of  children.  He  regarded  the  parasite  as  a  Cercomonad  and  termed 
it  Cercomonas  intestinalis^  which  name  as  a  rule  is  applied  to  Cercomonas  houiinis^ 
Davaine,  although  Stein  had  already  pointed  out  the  difference  between  the  two  species* 
Grassi  (1879)  observed  this  species  first  in  mice  (calling  it  Diinorphus  muris),  and 
subsequently  in  human  beings  in  Upper  Italy  and  named  it  Megastoma  entericum. 
Biitschli  and  Blanchard  then  laid  stress  on  the  identity  of  this  species  with  Lambl's 
C.  intestinalis  (1859),  and  consequently  called  it  Megastoma  intestinale.  Later, 
Blanchard  drew  attention  to  the  circumstance  that  the  generic  name  Megastoma 
chosen  by  Grassi  had  already  been  used  four  times  for  various  kinds  of  animals, 
and  established  the  genus  Lamblia.  Accordingly,  Z.  intestinalis  is  the  valid  name,, 
and  should  be  generally  adopted. 

In  Upper  Italy  the  parasite  in  the  encysted  condition  has  also  been  seen  by 
Perroncito  in  man.  At  the  same  time,  Grassi  and  Schewiakofif  began  a  new 
investigation  of  specimens  from  mice  and  rats.  In  Germany,  L.  intestinalis 
was  found  by  Moritz  and  Holzl,  Roos,  Schuberg  and  Salomon.  Moritz  and  Holzl 
confirmed  the  relative  frequency  of  the  species.  In  Konigsberg,  Prussia,  a  student 
found  encysted  La77iblia  in  his  fasces.  One  case  was  reported  from  Finland  by 
Sievers,  another  case  from  Scandinavia  by  Miiller.  Frshezjesski  and  Ucke  reported 
cases  from  Russia.  Jaksch  announced  the  occurrence  of  the  parasite  in  Austria  ; 
Piccardi  mentioned  their  presence  again  in  Italy.  They  were  reported  from  Egypt 
by  Kruse  and  Pasquale,  and  from  North  America  (Baltimore)  by  Stiles.  Noc  stated 
that  50  per  cent,  of  the  population  of  Tonkin  harboured  Lamblia.  Finally,  the 
structure  of  L.  intestinalis  has  been  described  by  Metzner  (1901),  and  by  Wenyon^ 
(1907)  in  mice. 

In  all  these  cases  L.  intestinalis  has  been  observed  in  the  small 
intestine,  or  in  the  evacuations  of  patients  with  intestinal  diseases. 
It  has  also  been  found  in  the  intestine  of  healthy  subjects.  Just  as 
Trichomonas  intestinalis  may  be  found  inhabiting  the  stomach  in 
diseases  of  that  organ,  in  which  an  alkaline  reaction  is  present 
(carcinoma),  so  has  L.  intestinalis  been  found  to  occur  under 
similar  circumstances  (Cohnheim,  Zabel).  However,  in  Schmidt's 
case,  I  per  cent,  hydrochloric  acid  was  certainly  stated  to  be  present. 
Infection  takes  place  by  the  ingestion  of  cysts  (fig.  20,  e),  as  was 
established  by  Grassi,  experimentally  on  himself.  Cereal  food-stuffs,, 
contaminated  with  Lamblia  cysts  from  vermin  of  the  locality,  such  as 
rats  and  mice,  serve  to  convey  the  infection  to  man.  Such  cysts  may 
probably  be  found  in  street-dust,  etc.  Stiles  induced  infection  in 
guinea-pigs,  and  Perroncito  in  mice  and  rabbits,  by  means  of  cysts 
of  Lamblia  from  human  beings.  Stiles  suspected  that  flies  could 
transport  Lamblia  cysts.  Mathis  (1914)  found  that  L.  intestinalis 
was  not  amenable  to  emetine,  at  any  rate  in  its  cystic  stage. 

Order.     Protomonadina,  Blochmann. 

The  smallness  of  the  Protomonadines  and  their  less  superficial 
situation  than  the  Polymastigines,  may  be  the  cause  that  so  far  as 
the  species  occurring  in  man  are  concerned,  they  were  formerly  less 
well  known.     As  regards  parasitic  species,  this  group  may  be  divided 

'  Arch.  f.  Protistenkunde,  Suppl.  i,  p.  169. 


CERCOMONAS   HOMINIS 


6i 


as  follows,  according  to  the  number  of  fiagella  and  the  presence  or 
absence  of  an  undulating  membrane  : — 

(i)  Cercomonadidce,  with  one  flagellum  at  the  anterior  extremity, 
without  an  undulating  membrane. 

(2)  Bodonidcv,  with  two  flagella,  without  an  undulating  membrane, 
except  in  Trypanoplasma. 

(3)  Tyypanosomidce,  with  one  flagellum,  and  an  undulating  mem- 
brane along  the  length  of  the  body  in  some  genera. 

Family.     Cercomonadidae,  Kent  emend.  Biitschli. 
Small  uniflagellate  forms,  without  cytostome. 

Genus.     Cercomonas,  Dujardin  emend.  Biitschli. 

Oval  or  rounded  organisms,  with  the  aflagellar  end  often  drawn 
out  into  a  tail-like  process. 

Cercomonas  hominis,  Davaine,  1854. 

Davaine  found  flagellates  in  the  dejecta  of  cholera  patients.  They 
had  pear-shaped  bodies,  lengthening  to  a  point  posteriorly.  Their 
length  was  from  10  /x  to  12  /x,  and  a  flagellum  about  twice  as  long  as 
the  body  projected  from  one  extremity  (fig.  21).  A  nucleus  was 
hardly  recognizable.  Occasionally  a  somewhat  long  structure  (cyto- 
stome ?)  appeared  at  the  anterior  extremity.  The  animals  moved  with 
remarkable  activity.  They  also  attached  themselves  by  means  of  their 
posterior  extremities  and  swung  about  around  the  point  of  attachment. 
Davaine  found  a  smaller  variety,  only  about  8  ix  long,  in  the  dejecta 
of  a  typhoid  patient  (fig.  21,  b). 


^-^ 


Fig.   21. — Cercomonas   hominis,  Dav.       a,  larger, 
bf  smaller  variety.    Enlarged.    (After  Davaine.) 


Fig.  22.  —  Cercomonas  hominis, 
Dav.  From  an  Echinococcus 
cyst.     (After  Lambl.) 


The  Flagellata  observed  by  Ekeckrantz  (1869)  in  the  intestine  of  man  belong  to 
this  form — at  least  to  the  larger  variety — and  Tham  (1870)  reported  fresh  cases  soon 
after.  Lambl's  publication  of  1875,  which  was  written  in  Russian,  and  became  known 
through  Leuckart's  work  on  parasites,  also  alludes  to  apparently  typical  Cercomonads, 
which,  however,  were  discovered,  not  in  the  intestine,  but  in  an  Echinococcus  cyst  in 
the  liver  (fig.  22).  The  elliptical,  fusiform,  rarely  pear-shaped  or  cylindrical  bodies 
of  the  parasites  measured  5  ^^  to  14  /*  in  length,  and  were  provided  with  a  flagellum 
at  one  end,  while  the  other  extremity  usually  terminated  in  a  long  point.  An  oral 
aperture  occurred  at  the  base  of  the  flagellum,  and  there  were  one  or  two  vacuoles 
near  the  posterior  extremity.     Longitudinal  division  was  also  observed  (fig.  22). 


62 


THE   ANIMAL   PARASITES   OF   MAN 


As  already  mentioned,  this  form,  which  Lambl  termed  Cerconionas 
intestinalis,  differs  considerably  from  the  form  found  by  the  same 
author  in  1859,  which  received  the  same  designation  {cf.  Lamhlia  intes- 
tinalis, p.  60),  but  it  corresponds  with  Cerconionas  hominis,  Davaine» 
The  latter,  as  well  as  C.  intestinalis,  Lambl,  1875,  is  usually  classed 
with  the  Trichomonads,  but,  as  has  already  been  remarked  {cf. 
Trichomonas  intestinalis,  p.  54),  this  cannot  be  considered  correct,  as 
only  one  flagellum  is  present. 

Cerconionas  vaginalis  (Castellani  and  Chalmers,  1909)  was  found 
in  the  vagina  of  native  women  in  Ceylon. 

Other  species  of  Cerconionas  have,  at  various  times,  been  recorded 
from  man.  However,  the  parasitic  species  of  the  genus  Cerconionas 
require  further  investigation. 


According  to  Janowski  (1896-7),  typical  Cercomonads  have  also  been  observed  in 
the  intestine  of  man  by  Escherich,  also  by  Cahen,  Massiutin,  Fenoglio,  Councilman  and 
Lafleur,  Dock,  Kruse  and  Pasquale,  Zunker,  Quincke  and  Roos,  and  others.  However, 
it  is  an  open  question  whether  the  Flagellata  observed  by  Roos  in  one  of  his  cases 
belonged  to  Davaine's  species,  the  size  showing  some  deviation  (14  jx  to  16  fx).  In 
his,  as  in  many  other  cases,  doubts  have  been  raised  as  to  whether  the  flagellates 
found  in  the  stools  had  actually  lived  in  the  intestine,  or  had  subsequently  appeared 
in  the  faeces :  for  this  a  surprisingly  short  time  only  is  necessary.  Salomon  also 
appears  to  have  observed  Cercomonads  {Berl.  klin.  Wochenschr.^  1899,  No.  46). 

As  with  T,  intestinalis  so  with  C.  ho?mms^  it  appears 
that  the  parasite  settles  not  only  in  the  intestine  but  also  in 
the  air-passages.  This  is  demonstrated  by  the  statements  of 
Kannenberg  and  Streng  of  the  occurrence  of  Monads  and 
Cercomonads  in  the  sputum  and  putrid  expectoration  in 
gangrene  of  the  lungs,  which  no  doubt  apply  to  C.  hominis 
{cf.  also  Artault).  Possibly  also  the  Flagellata  observed  in  the 
pleural  exudation  by  Litten  and  Roos  may  be  included  here  ; 
this  is  the  more  probable  in  Roos's  case  as  the  process  ensued 
in  the  pleura  after  the  breaking  through  of  a  vomica. 

Perroncito  and  Piccardi  have  described  encysted  stages  of 
Cercomonads. 


Monas  pyophila,  R.  Blanch.,  1895. 

Fig    2^  ^  Monas  ^*  ^^^"c^^''^   thus   designates   a   Flagellate   that    Grimm 

pyophila,  R.  Blanch.      found  in  the  sputum,  as  well  as  in  the  pus  of  a  pulmonary  and 
(After  Grimm.)  hepatic    abscess,  in  the  case  of  a  Japanese  woman  living  in 

Sapporo.  The  parasites  resemble  large  spermatozoa  (fig.  23). 
The  body,  30  ix  to  60 /x,  has  the  shape  of  a  heart  or  a  myrtle  leaf,  and  is  surrounded  by  a 
thick  cuticle  which  is  supposed  to  extend  into  the  interior  of  the  body,  dividing  it  into 
three  parts.  A  long  appendix  at  the  rounded  pole  is  covered  for  the  greater  part  of 
its  length  by  the  cuticle  ;  the  extremity,  however,  is  free  and  resembles  a  flagellum. 
The  parasites  were  very  active,  frequently  changed  their  shape,  and  were  able  to 
retract  the  long  appendix  within  the  body,  which  then  assumed  a  round  form. 
[This  organism  requires  further  investigation.] 


PROWAZEKIA   URINARIA  63 

Family.     Bodonidae,   Biitschli. 

Prolomonadina  which  are  either  free-Hving  or  parasitic,  with  two 
dissimilar  flagella,  while  the  possession  of  an  undulating  membrane 
and  of  a  kinetic  nucleus  or  blepharoplast  is  variable. 

There  are  three  genera  : — 

1.  Bodo,  Stein,   1878,  without  a  kinetic  nucleus  and  undulating 

membrane. 

2.  Prowazekia,     Hartmann    and    Chagas,    1910,    with    a    kinetic 

nucleus  and  without  an  undulating  membrane. 

3.  Trypanoplasma,   Laveran   and    Mesnil,    190 1,  with    a    kinetic 

nucleus  and  undulating  membrane. 
Of  these  genera  Prowazekia  must  be  discussed.  Bodo  does  not 
occur  in  man.  Species  of  Trypanoplasma  occur  in  the  blood  and  in 
the  gut  of  various  fishes,  in  the  seminal  receptacle  of  certain  snails, 
in  the  gut  and  genitalia  of  a  flatworm  (Dendrocoelum  ladeiim)  and  in 
the  vagina  of  a  leech.  Closely  allied  to  Trypanoplasma  is  the  genus 
Trypanophis,  parasitic  in  the  coelenteric  cavity  of  Siphonophores. 

Genus.     Prowazekia,  Hartmann  and  Chagas,  1910. 

The  genus  was  founded  for  a  flagellate  parasite,  Prowazekia  cruzi^ 
discovered  in  a  culture  of  human  faeces  in  Brazil.  Various  other 
species  have  been  referred  thereto.  The  genus  is  separated  from 
Bodo  by  the  possession  of  a  second  nucleus,  the  so-called  kinetonucleus 
or  blepharoplast.  It  differs  from  Trypanoplasma  in  the  absence  of  an 
undulating  membrane.  It  is  heteromastigote,  that  is,  it  possesses  two 
dissimilar  flagella,  one  anteriorly  directed  and  the  other  lateral  and 
trailing. 

The  principal  species  are  : 

Prowazekia  urinaria,  Hassall,  1859. 

Syn.  :   Bodo  urinartus^  Hassall,  1859;   Trichomonas  irregularis,  Salisbury,  1868; 
Cystomonas  urinaria,  Blanchard,  1885  ;  Plagiomonas  urinaria,  Braun,  1895. 

HassalP  in  1859  first  found  Bodo-like  flagellates  in  human  urine. 
He  examined  fifty  samples  of  urine  from  patients  suffering  from 
albuminuria  and  from  cholera.  The  reaction  of  the  urine  was  alkaline 
or  sometimes  only  feebly  acid.  The  flagellates  were  only  seen  after 
the  urine  had  been  standing  for  several  days.  Hassall  named  the 
organism  Bodo  urinarius,  and  gave  a  very  good  description  of  it  with 
illustrations.  The  flagellate,  which  was  round  or  oval,  measured  14  fi 
by  8  /jl.     The  organism  had  *^  one,  usually  two,  and  sometimes  three 

1  Lancet,  1859,  ii,  p.  503. 


64 


THE   ANIMAL   PARASITES   OF  MAN 


lashes  or  cilia."  In  1868  Salisbury  described  a  similar  flagellate  in 
the  urine  under  the  name  Trichomonas  irregularis.  Kiinstler  in  1883 
described  the  latter  parasite  under  the  name  B.  urinarius.  In  1885 
Blanchard,  considering  Kiinstler's  organism  a  different  parasite  from 
Hassall's,  called  it  Cystomonas  urinaria.  Braun,  in  1895,  gave  the 
name  Plagiomonas  urinaria.  Barrois  (1894)  considered  Kiinstler's 
and  Hassall's  organisms  to  be  identical  and  not  to  be  true  parasites 
of  man.  Sinton/  in  191 2,  found  the  flagellate  in  the  deposit,  after 
centrifuging,  of  a  24-hour  old  specimen  of  alkaline  urine  from  a 
Mexican  sailor  in  the  Royal  Southern  Hospital,  Liverpool.  Sinton 
found  a  kinetic  nucleus  or  blepharoplast  in  the  organism,  and 
therefore  placed  it  in  the  genus  Prowazekia. 


Fig.  24. — Types  of  Prowazekia  urinaria,     (a)  sausage-shaped  ;  {b)  round  ;  (<r)  carrot-shaped 

form.     (After  Sinton.) 

The  flagellate  stage  (fig.  24)  of  the  organism  is  polymorphic,  and  may 
be  either  {a)  sausage-shaped,  lo/x,  to  25  yit  in  length  by  2*5  fi  to  6  fi  in 
breadth  ;  (d)  round  or  oval,  varying  from  4  yit  in  diameter  to  oval 
forms  i^ /JL  by  10  fi;  (c)  a  carrot-shaped  form,  of  varying  size  up  to 
25  /x  by  4//,.  The  kinetic  nucleus  is  large  and  pear-shaped.  Near  it 
are  basal  granules,  closely  applied  to  one  another,  from  which  the 
flagella  arise.  There  is  a  small  cytostome  near  the  roots  of  the 
flagella.  There  is  a  well-marked  karyosome  in  the  nucleus.  The 
movement  is  jerky.  The  shorter,  anterior  flagellum  may  be  used  in 
food-capture.  In  life,  bacteria  have  been  seen  to  be  ingested.  Food- 
vacuoles  tend  to  accumulate  at  the  posterior  (aflagellar)  end.  A 
contractile  vacuole  may  be  present,  near  the  base  of  the  cytostome, 
and  may  really  be  the  dilated  fundus  of  the  latter.     Division  occurs  by 


Annals  Trop.  Med.  and  Parasitology,  vi,  p.  245. 


PROWAZEKIA   ASIATICA 


6s 


binary  fission.  The  organism  can  encyst  (fig.  25,  a)y  when  the  flagella  are 
lost,  and  round  or  oval  cysts  are  found,  5 //<  to  7  yu-  in  diameter.  After 
a  time  flagella  are  formed  inside  the  cyst,  and  the  organism  emerges 
therefrom  in  its  typical  flagellate  form  (fig.  25,  h — /). 

Sinton's  case  is  interesting.  He  obtained  the  flagellate  only  twice 
from  the  same  patient,  a  Mexican  then  in  hospital  in  Liverpool.  The 
flagellate  was  not  found  in  the  patient's  faeces,  nor  was  it  found  in 
the  urine  on  later  occasions  when  taken  aseptically. 


Fig.  25. — Proivazekia  urinaria.    Flagellate  emerging  from  cyst.     (After  Sinton.) 

In  cultures  Prowazekia  urmaria  was  always  found  in  association 
with  bacteria.  The  cultures  died  at  a  temperature  of  37°  C,  but  grew 
well  at  20°  C.  Various  media  were  useful  at  the  lower  temperature, 
such  as  urine,  salt  agar,  nutrient  agar,  serum  agar,  blood  agar,  peptone 
salt  solution,  and  diluted  blood  serum.  The  flagellate  was,  then, 
considered  to  be  an  accidental  contamination  and  not  a  true  parasite 
of  human  urine. 

Prowazekia  asiatica,  Castellani  and  Chalmers,  1910. 

The  flagellate  was  found  by  the  discoverers  in  the  stools  of  patients 
suffering  from  ankylostomiasis  and  diarrhoea  in  Ceylon.  It  was  referred 
by  them  to  the  gQnus  Bodo,  but  in  19 11  Whitmore^  further  studied 
it  and  placed  it  in  the  genus  Prowazekia.  In  the  stools  the  flagellate 
is  found  either  as  a  long,  slender  form  measuring  lOyLt  to  16  yu,  by  5  yu, 
to  Sjjb  or  as  a  rounded  form  S  /n  to  lo//,  in  diameter.  Its  cytoplasm  is 
alveolar.      A  rhizoplast    connects    the    basal   granules   to  the  kinetic 


Jrck.  f.  Protistenk.  xxii,  p.  370. 


66  THE   ANIMAL   PARASITES   OF   MAN 

nucleus.  There  is  multiplication  and  cyst  formation  as  before.  The 
organism  is  easily  cultivated,  especially  in  the  condensation  water  of 
nutrose  agar  and  maltose  agar.     The  pathogenicity  is  stated  to  be  nil. 

Prowazekia  javanensis,  Flu,  1912. 

Found  in  agar  cultures  from  the  motions  of  patients  at  Welte- 
vreden,  Dutch  East  Indies.'  The  flagellates  are  12  yu,  long  and  5  fi  broad. 
The  lateral  flagellum  is  stated  to  be  attached  to  the  cell  body  for  a 
short  distance.  Regarding  the  karyosome  in  the  nucleus,  the  author 
states  that  the  smaller  the  karyosome  the  more  chromatin  is  deposited 
on  the  nuclear  membrane.  Flu  mentions  that  the  specific  name 
javanensis  is  a  temporary  one,  as  in  the  course  of  time  it  may  be 
shown  that  there  is  only  one  species  of  Prowazekia. 

Prowazekia  cruzi,  Hartmann  and  Chagas,  1910. 

Found  in  a  culture  from  human  faeces  on  an  agar  plate  in  Brazil, 
and  considered  to  be  a  free-living  form.^  The  organism  is  oval  or 
pear-shaped,  8  yit  to  12  //,  long  and  5  /i,  to  6  /x  broad.  In  human  stools 
at  Tsingtau,  China,  a  Prowazekia  has  been  found  by  Martini  which 
he  thinks  is  the  same  as  Prowazekia  cruzi.  He  considers  it  to  be  a 
cause  of  human  diarrhoea  and  intestinal  catarrh. 

Prowazekia  Weinberg!,  Mathis  and  Leger,  1910. 

This  species  was  found  in  the  faeces  of  men,  both  healthy  and 
diarrhoeic,  in  Tonkin."  It  is  pear-shaped,  8/^  to  15  fju  long  by  4  yu,  to 
6*5  /t  broad.     The  flagella  occur  at  the  broad  end. 

The  discoverers  think  that  Prowazekia  weinbergi  is  an  intestinal 
inhabitant,  but  non-pathogenic,  since  it  was  found  to  occur  in  the 
faeces  even  when  obtained  with  aseptic  precautions. 

Prowazekia  parva,  Nagler,  1910. 

A  free-living  form  found  in  the  slime  on  the  stones  at  the  biological 
station  at  Lunz.  Another  Prowazekia  was  found  in  1914  in  tap-water 
in  Calcutta. 

Family.     Trypanosomidae,    Doflein. 

The  Trypanosomidae,  broadly  considered,  are  uniflagellate 
organisms,  the  flagellum  being  at  the  anterior  end.  The  flagellum 
arises  near  the  blepharoplast  (kinetic  nucleus),  which  lies  anterior, 
near  or  posterior  to  the  nucleus. 

'  Geneesk.  Tijdschr.  v.   Nederl.  Ind.,  lii,   p.  659;    Med.  v.  d.  Burg.   Geneesk.  d.  Nederl. 
Ind.y  iii,  p.  I. 

2  Mem.  Inst.  Osw.  Cruz.,  ii,  p.  64. 

^  Bull .  Soc.  Med.  Chir.  Indo-Chine,  i,  p.  471, 


TRYPANOSOMA  67 

The  following  genera  will  be  considered  : — 

Trypanosoma — with  an  undulating  membrane  along  the  length 
of  the  body. 

Crithidia — with  a  less  well-developed  undulating  membrane 
anteriorly  (see  fig.  49). 

Herpetomonas — including  the  so-called  Leptonionas,  with  anterior 
free  flagellum  only,  and  no  undulating  membrane. 

Lelshmania—non-{i3.ge\\3.te  forms  in  mammalian  blood,  flagel- 
late herpetomonad  stages  in  culture,  probably  occurring 
naturally  in  Arthropods. 

Genus.     Trypanosoma,   Gruby,   1843. 

The  members  of  the  genus  possess  a  single  fiagellum,  which 
arises  posteriorly,  adjacent  to  a  blepharoplast  or  kinetic  nucleus.  The 
flagellum  forms  a  margin  to  an  undulating  membrane,  and  may 
or  may  not  be  continued  beyond  the  body  as  a  free  flagellum. 
Many  species  are  parasitic  in  vertebrate  blood  and  in  the  digestive 
tracts  of  insects. 

Historical. 

The  history  of  blood  flagellates  goes  back  to  the  year  1841,  in  which  Valentin 
discovered  in  the  blood  of  a  brook-trout  {Salmo  fario  L.)  minute  bodies,  from  //i 
to  13 /x  in  length,  with  active  movements  and  presenting  marked  changes  in  form. 
Valentin  considered  the  parasite  a  new  species  of  the  old  genus  Proteus  or 
Amceba,  Ehrbg.  This  announcement  led  Gluge  (1842)  to  publish  a  similar  discovery 
he  had  made  in  frog's  blood.  The  latter  forms  were  called  by  Mayer  (1843)  Ainceba 
rotatoria^  Para7ncBciuin  loricatian  and  P.  costaiu7n^  while  Gruby  (1843)  called  them 
TryPa?iosorna  sanouinis}  Later  it  was  discovered  that  similar  organisms  occurred 
also  in  the  blood  of  birds  (Wedl  (1850),  Danilewsky)  and  of  mammals.  Gros  (1845) 
found  them  in  the  mouse  and  mole,  Chaussat  (1850)  in  the  house  rat,  Lewis  (1879)  in 
the  Indian  rat,  Wittich  (1881)  in  the  hamster.  Danilewsky  (1886-89)  and  Chalach- 
nikow  (1888)  investigated  the  structure  and  division  of  trypanosomes. 

In  the  case  of  all  these  forms,  there  was  no  discussion  as  to  a  pathogenic 
influence  on  the  host.  Opinion,  however,  as  to  the  action  of  trypanosomes 
changed  when,  in  1880,  Evans  found  flagellates  in  the  blood  of  horses  in  India 
that  suff'ered  from  a  disease  endemic  there  called  "  surra,"  and  associated 
the  parasites  with  the  disease.  Steel  and  Evans  were  successful  in  transmitting 
the  parasites — first  known  as  Spirochceta  evansj\  Steel,  then  as  Trichomonas 
evansi^  Crookshank,  and  finally  as  Trypanosoma  evansi — to  dogs,  mules  and  horses. 
They  recognized  that  the  above  mentioned  flagellates  in  the  blood  of  the  experi- 
mental animals  were  the  causal  agents  of  the  disease. 

From  that  time  there  was  a  considerable  increase  in  the  literature,  the 
contents  of  which  have  been  summarized  by  Laveran  and  Blanchard.  In  1894 
Rouget    discovered    trypanosomes    in    the    blood    of    African    horses    that    suffer 

1  Gruby's  generic  name  is  generally  accepted.  Still  others  have  been  used,  e.g.,  Undulinay 
Ray  I^ankester,  Globularia  Wedl,  Paramecioides  Grassi,  Trypanomonas  Danilewsky, 
Hcematotnonas  Mitrophanow. 


68  THE   ANIMAL   PARASITES   OF   MAN 

from  "  stallion's  disease  "  (dourine).  In  1894  Bruce  found  similar  forms  {T.  brucei)  in 
the  blood  of  South  African  mammals  suffering  from  "  nagana,"  and  in  consequence 
attention  was  drawn  to  the  part  which  the  much  dreaded  tsetse-fly  played  in 
the  transmission  of  "nagana."  In  1901  Elmassian  discovered  trypanosomes  in 
the  blood  of  horses  that  were  stricken  with  "  mal  de  caderas,"  which  is  very 
common  in  the  Argentine.  The  disease  in  cattle  named  "galziekte"  (gall- 
sickness),  occurring  in  the  Transvaal,  was  also  at  one  time  attributed  to 
a  trypanosome  remarkable  for  its  great  size,  and  like  some  other  species,  bearing 
the  name  of  its  discoverer  {T.  theileri). 

The  study  of  the  species  hitherto  known  has  been  carried  on  partly  by  the 
above  mentioned  authors  and  in  part  by  others,  e.g.^  Rabinowitsch  and  Kempner, 
Laveran  and  Mesnil,  Wasiliewski,  Senn.  It  was  greatly  advanced  by  the  method 
of  double  staining  (with  alkaline  methylene  blue  and  eosin)  introduced  by 
Romanovvsky  (1891)  and  elaborated  by  Ziemann,  Leishman,  Giemsa  and  others. 
By  this  means  the  presence  of  a  terminal  flagellum  and  of  an  undulating  niembrane 
at  the  side  of  the  flattened  and  extended  body  was  demonstrated.  Laveran  and 
Mesnil  (1901)  discovered  allied  flagellates  in  the  blood  of  the  fish,  Scardiniiis 
erythrophthalmus.  These  flagellates,  now  placed  in  the  genus  Trypanoplasma,  had 
a  second  free  flagellum  in  addition  to  the  one  bordering  the  undulating  membrane. 
Trypanoplasms  have  since  been  found  in  both  freshwater  and  marine  fishes.  The 
transmission  of  trypanoplasms  of  freshwater  fishes  is  effected  by  leeches.  Trypano- 
plasma  varium  from  Cobitis  is  transmitted  by  Heniiclepsis  viarginata  according  to 
Leger,  while  the  Trypanoplasmata  of  Cypriniis  carpio  and  Abramis  bravia  reach 
new  hosts  by  the  agency  of  Piscicola  according  to  Keysselitz. 

Another  ally  of  the  Trypanosomidae,  Trypanophis^  lives  in  the  ccelenleric 
cavity  of  Siphonophores.  It  has  also  an  extra  terminal  flagellum  (Poche, 
Keysselitz).     [^Trypanoplasma  and  Trypanophis  belong  to  the  Bodonidce^  see  p.  63]. 

Finally  it  was  shown  that  Trypanosomes  occurred  in  human  beings.  Although 
Nepveu's  early  report  of  trypanosomes  in  the  blood  of  malarial  patients  may  be 
doubtful,  subsequent  researches  by  Forde  and  Button  demonstrated  trypanosomes 
(fig.  28)  in  the  blood  of  a  European,  apparently  suffering  from  malaria,  living  in  the 
Gambia.  Button  (1902)  called  the  human  trypanosome,  T.  gambiense.  The  expedi- 
tion despatched  by  the  Liverpool  School  of  Tropical  Medicine  (1902)  to  Senegambia 
found  trypanosome  infections  in  six  cases  among  a  thousand  inhabitants  examined. 

About  the  same  time  attention  was  devoted  to  the  disease  of  West  African 
negroes  known  for  a  century  as  "  sleeping  sickness."  Castellani  (1903)  was  the 
first  to  succeed  in  demonstrating  the  presence  of  trypanosomes  (at  first  called 
T.  ugandense)  in  centrifugalized  cerebro-spinal  fluid  obtained  by  puncture  from  cases 
of  sleeping  sickness  in  Uganda.  Similar  discoveries  were  made  by  Bruce,  who  also 
found  trypanosomes  in  the  blood  of  those  attacked  with  sleeping  sickness.  Sambon 
regarded  a  species  of  Glossina  as  the  transmitter.  From  consideration  of  the 
geographical  distribution  of  the  disease  Christy  regarded  Glossina paipalis  as  the 
transmitter.  Brumpt  first  thought  it  was  G,  morsitans^  but,  later,  supported  the 
view  of  G.  paipalis.  Bruce,  Nabarro  and  Greig  also  named  the  same  insect  as 
the  transmitter,  not  only  for  geographical  reasons  but  also  because  healthy  apes 
became  infected  by  the  bite  of  certain  G.  paipalis.  The  inoculation  of  cerebro- 
spinal fluid  from  subjects  of  sleeping  sickness  into  the  spinal  canal  of  apes  {Macacus) 
had  the  same  result. 

Just  as  the  discovery  of  the  malarial  parasites  called  forth  a  whole  flood 
of  research  memoirs  which  were  followed  by  a  second  series  on  the  relation  of 
the  mosquitoes  to  malaria,  so  a  similar  outpouring  occurred  after  the  discovery 
of  the  pathogenic  trypanosomes  of  mammals  and  men.  In  both  cases  the  inquiry 
was   not   limited   to   the   stages   in    man   and  other  vertebrate   hosts,  but  the  fate 


TRYPANOSOMES  69 

of  the  parasites  in  the  intermediate  (invertebrate)  hosts  was  investigated,  and 
allied  species   were  obtained  from   many  different  hosts. 

Novy  and  MacNeal  (1903)  were  the  first  to  cultivate  trypanosomes  in  artificial 
media  (blood-agar). 

In  1910  Stephens  and  Fantham  recorded  the  presence  of  another  human 
trypanosome,  T.  rhodesiense,  from  a  case  of  sleeping  sickness  in  Rhodesia,  where 
G.  falpalis  was  absent.  Kinghorn  has  since  demonstrated  that  T.  rhodesiense  is 
transmitted  by  G.  inorsita?is,  Kinghorn  and  Yorke  believe  that  big  game  {e.g.^ 
antelope)  is  the  reservoir  of  T.  rhodesiense. 

The  output  of  literature  on  trypanosomiasis  in  men  and  animals  is  enormous. 
To  cope  with  it  the  Sleeping  Sick?iess  Bureau  Bulletin  was  founded  in  1908, 
and  it  is  now  (since  November  191 2)  continued  as  a  section  of  the  Tropical  Diseases 
Bulletin^  wherein  current  literature  is  reviewed. 

General. 

Trypanosomes  occur  in  the  blood  of  representatives  of  all  the 
vertebrate  classes.  Often  the  trypanosomes  occur  so  scantily  in  the 
blood  that  they  are  overlooked  on  examination.  A  useful  aid  in 
detecting  the  flagellates  in  such  cases  consists  in  the  use  of  cultures 
of  the  blood  of  the  host  on  artificial  media.  Stimulated  by  the 
medium  multiplication  occurs,  and  hence  the  parasites  are  more 
easily  detected.  [For  the  composition  of  such  culture  media  see 
Appendix.] 

There  is  a  periodicity  in  the  appearance  of  the  trypanosomes  in 
the  peripheral  blood  of  the  host,  due  to  alternating  phases  of  multi- 
plication and  of  rest  on  the  part  of  the  parasites.  Such  periodicity 
has  been  established  both  by  biological  and  enumerative  methods. 
Again,  a  seasonal  variation  has  been  observed  in  the  occurrence 
of  certain  trypanosomes  in  the  peripheral  circulation  of  the  hosts  ;  for 
example,  some  trypanosomes  {e,g.^  T.  noctiice  in  birds)  are  found  only 
in  the  summer  in  the  blood,  while  in  the  winter  they  occur  in  the 
internal  organs. 

Recent  cultural  researches  have  established  that  trypanosomes,  e.g., 
T.  mnericamnn,  may  be  present  in  very  small  numbers  in  hosts,  such  as 
cattle,  which  are  quite  unharmed  by  them,  and  in  which  the  presence 
of  these  flagellates  formerly  was  never  suspected  ("  cryptic  trypano- 
somiasis.") However,  the  majority  of  the  trypanosomes  occurring  in 
domestic  animals  are  usually  deleterious  or  even  lethal  to  their  hosts. 
Many  wild  animals,  such  as  various  species  of  antelope,  harbour 
trypanosomes  without  being  injured  thereby.  In  such  cases  it  is 
probable  that  the  vertebrate  hosts  have  been  so  long  parasitized  in  the 
past,  that  they  have  become  tolerant  and  immune  to  the  effects  of  the 
flagellates.  Should  such  trypanosomes  of  wild  animals  be  transmitted 
to  domesticated  stock  or  man,  they  may  re-acquire  their  initial  viru- 
lence and  become  pathogenic  to  the  new  host.  As  a  general  state- 
ment, the  newer  a  parasite  is  to  its  host  the  greater  is  its  virulence. 


70 


THE   ANIMAL   PARASITES   OF   MAN 


For  example,  T.  gmnbiense,  T.  rJiodesiense  a.nd  T.  bnicei  me  innocuous 
to  big  game  in  Africa,  but  are  pathogenic  to  man  and  domestic  animals 
respectively.  ,  Pathogenic  trypanosomes  appear  to  have  a  wider  range 
of  hosts,  that  is,  to  be  less  limited  to  one  specific  host  than  non- 
pathogenic forms.  Thus,  7\  rhodesiense  is  pathogenic  to  man  and  all 
laboratory  animals,  while  it  is  non-pathogenic  to  antelopes  and  their 
kind. 

Morphology. 

The  general  structure  of  the  various  trypanosomes  shows  much 
uniformity,  though  variations  in  size  and  shape  occur.  Typically  the 
body  is  elongate  and  sinuous..  The  flagellar  end  tapers  gradually  to 
a  point,  the  aflagellar  extremity  usually  being  rounded  or  more  blunt. 
In  some  trypanosomes  there  is  much  diversity  in  size,  the  organisms 
varying  from   long,  slender  forms  to   short,  stumpy  ones  ;    in  other 


Fig.  26. —  Trypanosovia  brucei  in  division.     ;;,  nucleus;  IjI,  blepharoplasl  ;  Jl,  flagellum. 
X  2,oco.     (After  Laveran  and  Mesnil.) 

species  relative  constancy  of  size  is  maintained^  The  former  are 
known  as  polymorphic  trypanosomes,  the  latter  as  monomorphic 
forms. 

Two  nuclei  are  present.  The  main  or  principal  nucleus,  some- 
times termed  the  trophic  nucleus,  is  often  situated  towards  the  centre 
of  the  body  ;  it  is  frequently  of  the  vesicular  type,  containing  a 
karyosome.  The  blepharoplast  or  kinetic  nucleus  is  posterior  to  the 
nucleus,  and  usually  is  rod-like.  The  flagellum  arises  close  to  the 
blepharoplast,  and  forms  an  edge  to  the  undulating  membrane. 
It  may  or  may  not  extend  beyond  the  limits  of  the  undulating 
membrane.  If  it  does  so,  the  unattached  part  is  known  as  the  free 
flagellum.  Sometimes  a  small  granule  is  found  at  the  origin  of  the 
flagellum.  This  is  the  basal  granule,  and  is  considered  by  some  to 
function  as  the  centriole  of  the  kinetic  nucleus. 


TRYPANOSOMES 


71 


The  undulating  membrane  is  a  lateral  extension  of  the  ectoplasm 
or  periplast,  and  is  the  maiji  agent  in  locomotion.  It  is  edged  by  the 
flagellum,  which  forms  a  deeply  stainable  border  to  it.  Within  the 
membrane  substance,  often  arranged  parallel  with  its  edge,  are  a 
number  of  fine  contractile  elements,  the  myonemes.  These  con- 
tractile elements  may  also  occur  on  the  body  of  the  trypanosome. 
They  are  easily  seen  in  some  large  trypanosomes,  but  are  difficult  of 
demonstration  in  others,  owing  to  their  great  fineness. 

Multiplication  of  trypanosomes  in  the  blood  is  brought  about  by 
binary  longitudinal  fission  (fig.  26).  Division  is  initiated  by  that  of 
the  blepharoplast  and  nucleus.  The  division  may  be  equal  or  sub- 
equal,  whereby  differences  in  size  of  individuals  partly  arise.     Multiple 


Fig.  27. —  Trypanosoma  leivisi.    Multiplication  rosettes.    X  i,ooo.    (After  Laveran  and  Mesnil.) 

division  by  repeated  binary  fission,  without  complete  separation  of 
the  daughter  forms,  is  known  in  some  trypanosomes  {e.g.y  T.  lewisi)^ 
and  rosettes  of  parasites  thereby  are  produced  (fig.  27). 

The  classification  of  trypanosomes  is  very  difficult.  Laveran 
(1911)^  has  suggested  the  examination  of  the  relative  length  of  the 
flagellum  as  a  diagnostic  character,  and  so  arranged  these  flagellates 
in  mammals  in  three  groups.  The  first  group  included  those  trypano- 
somes always  having  part  of  the  flagellum  free  {e.g.,  T.  evansiy 
T.  vivax)  ;    the   second    group    comprised    forms  without   a    part   of 

'  Ann.  Inst.  Pasteur,  xxv,  p.  497. 


72  THE   ANIMAL   PARASITES   OF   MAN 

the  flagellum  free  (e.g.,  T.  congolcnse),  while  the  third  group  included 
forms  some  members  of  which  have  free  flagella,  while  others  have 
not  {e.g.,  T.  gainhiense).  Bruce^  (1914)  ^^'^^  Yorke  and  Blacklock^ 
(1914)  have  also  devised  classifications. 

Resting  stages  of  some  trypanosomes  have  been  found  in  the 
mternal  organs  of  their  vertebrate  hosts.  The  formation  of  these 
oval,  Leishmania-like  bodies  will  be  noted  in  individual  cases  later. 
Similar  small  oval  bodies  form  an  important  phase  in  the  life-history 
of  T.  cruzi,  which  multiplies  normally  by  multiple  fission  or  schizogony 
into  these  oval,  daughter  elements,  and  not  by  binary  longitudinal 
fission  in  the  circulating  blood. 

Polymorphism  in  trypanosomes  (e.g.,  T.  gambiense,  T.  rhodesiense) 
is  now  interpreted  as  a  phenomenon  resulting  from  growth  and 
division.^  Long,  thin  forms  are  those  about  to  divide.  Fully  mature 
forms  are  shorter  and  broader.  Various  intermediate  types  occur  and 
represent  growth  forms.  Formerly,  polymorphism  was  interpreted  in 
terms  of  sex,  thin  forms  being  regarded  as  males,  broad  forms  as 
females,  while  the  intermediate  types  were  termed  indifferent.  Con- 
jugation was  not  observed,  and  there  is  no  evidence  in  support  of 
the  sexual  interpretation. 

The  transmission  of  trypanosomes  from  one  vertebrate  host  to 
another  is  usually  accomplished  by  the  intermediation  of  some  biting 
arthropod  in  the  case  of  terrestrial  animals,  while  leeches  are  usually 
considered  to  act  as  transmitters  in  the  case  of  the  trypanosomes 
occurring  in  aquatic  animals.  Developmental  phases  of  the  life- 
histories  of  trypanosomes  occur  in  the  invertebrate  transmitters,  and 
will  be  considered  in  individual  cases. 

Trypanosoma  gambiense,  Button,  1902. 

Syn.  :  Trypanosoma  hominis^  Manson,  1903.  Trypajtosoma  nepveui,  Sambon,  1903. 
Trypanosoma  castellanii,  Kruse,  1903.  Trypanosoma  ugandense,  Castellani, 
1903.     Trypanosoma  fordii^  Maxwell  Adams. 

In  vertebrate  blood  Trypanosoma  gambiense  is  polymorphic,  for 
long,  thin  forms  may  be  seen  in  contrast  with  short,  stumpy  forms, 
as  well  as  intermediate  forms  (fig.  29,  a — c).  This  polymorphism  has 
been  interpreted  in  terms  of  sex,  especially  by  German  investigators, 
following  Schaudinn  (see  above).  However,  there  is  no  evidence  of 
conjugation,  and  the  polymorphic  forms  are  more  easily  interpreted 
in  terms  of  growth  and  division,  for  the  long  thin  forms  are  potential 
dividing  organisms,  and  the  stumpy  or  short  parasites,  with  little  or 
no  free  flagellum,  are  the  adult  individuals. 

'  Trans.  Soc.  Irop.  Med.  (Sr"  Hyg.,  viii,  p.  i. 

^  Annals  Trop.  Med.  and  Parasitol.,  viii,  p.  I. 

'^  Robertson  (191 2),  Proc.  Roy.  Soc,  B,  Ixxxv,  p.  527. 


TRYPANOSOMA   GAMBIENSE 


73 


Morphology  of  T.  gamhietise  in  the  Circulaihig  Blood. 

T.  gamhiense  varies  from  13  fju  to  ^6  jul  in  length,  its  average  length 
being  24*8  yu,  as  was  determined  in  19 13  by  exact  biometrical  methods 
by  Stephens  and  Fantham.^  ThT^eeXorms^of^i^asite  occur.  According 
to  Miss  Robertson,^  the  relatively 
short  forms  from  13  yit  to  21  /z 
long  may  be  regarded  as  the 
matureor  *^adult"  type  of  parasite 
in  the  blood.  They  carry  on  the 
cycle  in   the   vertebrate.     From 


them  intermediate  forms,  which 
are  longer  than  the  ''adult"  but 
at  first  have  the  same  breadth, 
arise  by  growth.  They  possess 
a  free  flagellum.  The  intermedi- 
ate forms  grow  into  long  indi- 
viduals, which  are  those  about  to 
divide.  The  products  of  division  give  rise,  directly  or  indirectly,  to 
the  adult  forms. 

The  organism  has  an  elongate  body  with  an  anterior  or  flagellar 
end   and  a  blunter  posterior  or  non-flagellar  end.     The  protoplasm 


Trypanosoma  gambiense. 
(After  Button. ) 


X  1,700. 


Fig.  29.  —  Trypanosoma  gambiense.  Development  in  vertebrate  host,  a^  long,  slender, 
b,  intermediate  and  c,  short,  stumpy  forms,  found  in  the  blood  ;  d,  e,  /,  non-flagellale,  latent 
forms  from  internal  organs.      x  2,000.     (Original.     From  preparations  by  Fanlham.) 

is  finely  granular,  large  inclusions  being  rare.  The  central  nucleus  is 
oval  and  large,  often  containing  most  of  its  chromatin  concentrated  as 
a  karyosome,  with  small  granules  only  scattered  near  or  on  the  fine 
nuclear   membrane.      The    blepharoplast   is   either  rounded   or   rod- 


'  Annals  Trop.  Med.  and  ParasitoL^  vii,  p.  27.         -  Phil.  Trans.,  B  (1913),  cciii,  pp.  161-184. 


74  THE   ANIMAL   PARASITES   OF   MAN 

shaped.  The  undulating  membrane  is  thrown  into  folds  and  is 
bordered  by  the  fiagellum.  A  small  basal  granule  may  be  present 
near,  or  at  the  actual  origin  of  the  flagellum. 

Multiplication  in  the  vertebrate  is  brought  about  by  longitudinal 
division.  According  to  the  recent  account  of  division  by  Miss 
Robertson,  the  blepharoplast  doubles,  then  the  flagellum  splits  for  the 
greater  part  of  its  length,  and  the  daughter  flagella  separate,  one  being 
shorter  than  the  parent  flagellum.  The  nucleus  often  shows  two  well 
marked  dark  granules  on  ihe  membrane  at  opposite  poles,  and  these 
appear  to  act  as  centrosomes.  Nuclear  constriction  occurs  and  the 
halves  gradually  separate.  Finally  the  two  daughter  organisms  become 
free,  the  aflagellar  end  splitting  last.  The  products  of  division  may  be 
equal  or  unequal.  Repeated  division  goes  on  in  the  general  circulation 
until  the  blood  swarms  with  parasites.  Then  the  trypanosomes  gradually 
disappear,  and  a  period  occurs  when  it  is  practically  impossible  to 
demonstrate  the  parasite  in  the  blood.  At  such  a  period,  trypanosomes 
can  be  obtained  by  puncture  of  the  enlarged  lymphatic  glands  or  of 
the  spinal  canal,  or  can  be  found  in  the  internal  organs,  more  particu- 
larly in  the  spleen,  lungs,  liver  and  bone-marrow.  In  the  latter 
organs,  latent  bodies  are  produced  (fig.  29,  d — -/)  which  are  capable  of 
again  becoming  flagellates  and  entering  the  general  circulation. 
Their  formation  was  described  by  Fantham  (1911).^  The  parasite 
contracts,  the  blepharoplast  migrates  towards  the  nucleus,  a  very  thin 
coat  differentiates  around  the  two  nuclei  and  a  certain  amount  of  cyto- 
plasm, and  the  parts  exterior  to  the  coat  disintegrate,  leaving  a  small, 
oval  body  behind.  Fuller  details  are  given  in  connection  with 
T.  rhodesiense.  Laveran  (191 1)^  considers  that  latent  bodies  are 
'^  involution "  forms,  but  acknowledges  that  they  can  flagellate  and 
become  infective  in  fresh  blood. 

No  multiplication  of  the  trypanosomes  w^ithin  the  cells  of  the 
lung,  liver  or  spleen  of  infected  monkeys  was  found  by  Miss 
Robertson  in  her  recent  researches. 

There  appear  to  be  negative  periods  in  infected  monkeys,  since, 
although  trypanosomes  may  occur  in  their  blood  at  such  times,  they 
are  not  infective  to  Glossina. 

Development  in  Glossina  palpalis. — The  principal  accounts  are 
those  by  Sir  D.  Bruce  and  his  colleagues  (1911),^  and  by  Miss 
Robertson^  (1912),  whose  results  will  be  followed.  According  to  the 
latter  investigator  T.  gambiense  never  enters  the  body  cells  of  the  fly 
(G.  palpalis),  nor  does  it  penetrate  the  gut  wall  into  the  body  cavity. 
Practically  no  crithidial  stage  occurs  in  the  fly's  main  gut,  but  a 
trypanosome  facies  is  retained  therein. 

After  the  trypanosomes  are  ingested  by  the  fly  during  a  meal  of 

'  Froc.  Roy.  Soc.^  B,  Ixxxiii,  p.  212.  '^  C.  R.  Acad.  Set.,  153,  p.  649. 

^  Froc.  Roy.  Soc,  B,  Ixxxiii,  p.  513.  ^  Froc.  Roy.  Soc,  B,  Ixxxvi,  p.  66. 


TRYPANOSOMA   GAMBIENSE 


75 


infected  blood,  sooner  or  later  multiplication  occurs.  This  develop- 
ment usually  begins  in  the  middle  or  posterior  part  of  the  mid  gut, 
and  Irypanosomes  of  varying  sizes  are  produced.  After  the  tenth 
or  twelfth  day,  many  long,  slender  trypanosomes  (tig.  30,  a)  are 
found,  which  gradually  move  forwards  into  the  proventriculus.  Such 
long,  slender  forms  represent  the  limit  of  development  in  the  lumen 
of  the  main  gut.  The  proventricular  type,  developed  about  the 
eighth  to  the  eighteenth  or  twentieth  day,  is  not  infective ;  it  may 
occur  in  the  crop,  but  is  not  to  be  found  permanently  there. 
Between  the  tenth  and  the  fifteenth  days  multinucleate  forms  of 
trypanosomes  are  found,  and  may  be  styled  multiple  forms  (fig.  30,  b). 
Some  of  these  latter  may  be  degenerative. 


Fig.  30.  —  Trypanosoma  gambiense.  Development  in  the  fly,  Glossina  palpalis.  a,  slender, 
proventricular  form;  b,  multinucleale  form;  r,  d,  crithidial  forms;  ^,  infective  type  of 
trypanosonie  found  in  salivary  gland.      x  2,500.     (After  Robertson.) 

Invasion  of  the  Salivary  Glands  of  the  Fly. — Long,  slender  trypano- 
somes from  the  proventriculus  pass  forward  into  the  hypopharynx. 
They  then  pass  back  along  the  salivary  ducts,  about  sixteen  to  thirty 
days  after  the  fly's  feed.  The  trypanosomes  reach  the  salivary  glands 
_as  long,  slender  forms.  In  the  glands  they  become  shorter  and 
broader,  attach  themselves  to  the  surrounding  structures,  and  assume 
the  crithidial  facies  (fig.  30,  c,  d).  As  crithidial  forms  they  remain 
attached  to  the  wall  and  multiply  in  the  glands.  These  crithidial 
stages  differentiate  into  the  short,  broad  trypanosome  forms,  capable 
of  swimming  freely  (fig.  30,  e). 

Miss  Robertson  considers  the  development  in  the  main  gut  to  be 
indifferent  multiplication,  and  that  salivary  fluid  seems  necessary  to 
stimulate  trypanosomes  to  the  apparently  essential  reversion  to  the 


76  THE    ANIMAL    PARASITES    OF   MAN 

crithidial  type.  The  second  development  in  the  saUvary  gland  is  the 
essential  feature.  The  short,  stumpy  forms  of  trypanosomes  (fig.  30,  e) 
finally  produced  in  the  salivary  glands  are  alone  infective.  No 
conjugation  of  trypanosomes  occurs  in  the  fly.  Only  about  5  per 
cent,  of  captive  tsetse  flies  fed  on  trypanosome-infected  blood  become 
infective,  but  they  probably  remain  infective  for  the  rest  of  their  lives. 

J.  G.  Thomson  and  Sinton  (1912)^  have  obtained  in  cultures  the 
various  trypanosome  forms  of  T.  gamhiense  seen  in  the  fly's  main  gut. 

Duke  (1912)2  found  T.  gamhiense  in  a  species  of  antelope,  the 
situtunga  (Tragelaphus  spekei),  on  Damba  Island  in  Victoria  Nyanza. 
Wild  G.  palpalis  could  be  infected  therefrom.  The  antelope  may 
then  act  as  a  sleeping  sickness  reservoir  in  that  district,  but  men  are 
apparently  the  chief  reservoir. 

Trypanosoma  nigeriense,  Macfie,  1913.^ 

Macfie  has  recently  (August,  19 13)  described  a  human  trypano- 
some from  the  Eket  district  of  Southern  Nigeria.  It  is  common  in 
young  people.  The  disease  produced  does  not  seem  to  be  of  a 
virulent  type  in  Nigeria,  and  does  not  occur  in  epidemic  form.  In 
the  early  stages  the  glands  of  the  neck  are  enlarged.  'In  the  later 
stages — cases  of  which  are  rarer — lethargy  appears.  The  parasite 
is  a  polymorphic  trypanosome,  morphologically  almost  indistinguish- 
able from  T.  gamhiense,  though  it  may  be  slightly  shorter.  Macfie 
recorded  the  occurrence  in  his  preparations  of  a  few  trypano- 
somes appearing  to  have  a  flagellum  free  during  their  whole  length. 
Some  of  the  parasites,  as  seen  in  a  sub-inoculated  guinea-pig,  are  very 
small  (8  yu,  long).  Other  trypanosomes  have  their  nuclei  displaced 
somewhat  anteriorly.  This  parasite  may  only  be  a  variety  of 
T.  gamhiense.     The  parasite  is  perhaps  spread  by  Glossina  tachinoides, 

Trypanosoma  rhodesiense,  Stephens  and  Fantham,  1910. 

The  parasite  was  found  in  the  blood  of  a  young  Englishman  who 
had  contracted  sleeping  sickness  in  the  Luangwa  Valley,  North- 
eastern Rhodesia,  in  the  autumn  of  1909.  The  patient  had  never 
been  in  an  area  infested  with  Glossina  palpalis. 

(i)  Morphology. — The  morphology  of  the  parasite  in  man  and 
sub-inoculated  rats  was  studied  by  Stephens  and  Fantham  in  1910.* 
They  pointed  out  a  morphological  peculiarity  in  the  presence  of 
certain  trypanosomes  with  posterior  nuclei  in  sub-inoculated  animals, 
that  is,  parasites  in  which  the  nucleus  (trophonucleus)  was  situated 
towards  the  posterior  or  aflagellar  end,  close  up  to  or  even  beyond  the 

*  Annals  Trop.  Med:  and Parasitoh,  vi,  p.  331.     "  Proc.  Roy.  Soc,  B,  Ixxxv,  pp.  156,  483. 
'^Annals  Trop.  Med.  and  Parasitol.,  vii,  p.  339;  viii,  p.  379. 

*  Proc.  Roy.  Soc,  B,  Ixxxiii,  p.  28. 


TRYPANOSOMA   RHODESIENSE 


77 


blepharoplast  or  kinetic  nucleus  (fig.  31,4,  5).  When  the  nucleus  was 
besjde  the  blepharoplast,  the  former  was  seen  to  be  kidney-shaped 
(fig.  31,4).  The  posterior  nuclear  forms  were  of  the  stout  and  stumpy 
variety,  and  about  6  per  cent,  of  the  stumpy  forms  were  found  to  have 
their  nuclei  displaced  from  the  centre.  The  anterior  or  flagellar  end 
of  these  trypanosomes  often  contained  chromatoid  granules.  T. 
rhodesleiise  VRvies  in  length  from  12 //,  to  39 /a^;  short  stumpy  forms 
vary  from  13  fi  to  21  //,,  intermediate  forms  from  21  fM  to  24.  fi,  and  long, 
slender  forms  from  25  jjl  onwards.     The  average  length  is  24*1  /x. 

Certain  regular  periods  occur  in  the  course  of  the  trypanosomiasis 
when  few  or  no  flagellate  trypanosomes  are  found  in  the  peripheral 
blood  of  the  patient  or  of  the  sub-inoculated  animal.  These  periods 
can  be  explained  in  terms  of  morphology,  for  the  trypanosomes  are 


Fig.    31. — Trypauosoma   rkodesiense.       i,  Long  narrow   form;    2-4,   nucleus    passing   to 
posterior  (aflagellar)  end  ;  5,  nucleus  quite  posterior.    X  1,800.    (After  Stephens  and  Fantham.) 


capable  of  assuming  a  non-flagellate  form  in  the  internal  organs  of  the 
host,  particularly  in  the  lungs  and  in  the  spleen.  Such  forms  are 
known  as  ^'latent"  or  ''resting"  forms.  The  term  ''latent  body" 
was  first  used  by  Moore  and  Breinl  in  1907^  in  connection  with 
T.  gambiense.  Fantham^  (191 1)  has  described  the  process  of  forma- 
tion of  latent  from  motile  forms  and  the  reconversion  of  the  latent 
bodies  into  active  flagellates.  Fresh  preparations  of  splenic  blood 
or  lung  blood  containing  trypanosomes  were  made.  A  trypano- 
some  gradually  withdrew  or  cast  off  its  flagellum,  concentrated  its 
cytoplasm,  and  became  more  or  less  elongate  oval.  Nucleus  and 
blepharoplast  approached  one  another  and  came  to  lie  more  or  less 
side  by  side.  Then  an  opaque  line  often  made  its  appearance  around 
the  nuclear  area  and  differentiated  as  a  slight  envelope  or  covering, 
the  cytoplasm  external  to  this  merely  degenerating.  The  small,  oval, 
refractile  body  (fig.  29,  d—f)  thus  formed  was  a  non-flagellate  latent 
body,  2  /x  to  4  yLt  in  diameter,  like  Leishinania  or  the  non-flagellate, 

'  Stephens  and  Fantham  (1912-13),   Proc.  Roy.  Soc,  B,  Ixxxv,  p.  223,  and  Annals   Trap. 
Med.  and  Parasitol.,  vii,  p.  27, 

Annals  Trop.  Med.  and  Parasitol.,  i,  p.  441.         ^  Proc.  Roy.  Soc,  B,  Ixxxiii,  p.  212. 


78  THE   ANIMAL   PARASITES   OF   MAN 

multiplicative  forms  of  T.  cnizi  (fig.  34),  and  remains  temporarily 
inactive  in  the  internal  organs  of  the  host.  After  this  period  of 
inactivity,  the  non-flagellate  body,  recuperated  by  its  rest,  begins 
to  elongate  again.  The  nuclei  separate.  From  a  small  vacuole-like 
portion  the  fiagellum  differentiates  and  forces  out  the  ectoplasm, 
which  assumes  the  form  of  the  imdulating  membrane  with  its 
flagellar  border.  Subsequent  grow^th  results  in  the  production  of  the 
typical  trypanosome  form,  wiiich  re-enters  the  circulating  blood  and 
multiplies  by  longitudinal  binary  fission.  Division  of  the  parasite 
prior  to  the  formation  of  a  latent  body  may  occur  and  division 
of  the  latent  forms  themselves  is  known,  though  less  common. 
Consequently  latent  bodies,  like  the  flagellate  forms  themselves, 
show  diversity  in  size.  The  blepharoplast  of  the  latent  bodies  is 
sometimes  less  well  marked  than  in  Leishmania  (see  fig.  29,  d — -/). 
Laveran's  views  on  these  bodies  have  already  been  given  on  p.  74. 

(2)  Animal  Reactions. — The  posterior  nuclear  trypanosomes  were 
found  in  all  sub-inoculated  animals,  such  as  rats,  guinea-pigs,  dogs, 
mice,  Macacus,  rabbits  and  horses,  but  were  not  seen  in  the  human 
patient,  as  few  trypanosomes  occurred  in  his  peripheral  blood. 
R.  Ross  and  D.  Thomson^  found  a  periodic,  cyclical  variation  in  the 
number  of  the  parasites  in  the  patient's  blood  from  day  to  day,  the 
cyclical  period  being  about  a  week  (fig.  32).  Fantham  and  J.  G. 
Thomson^  (i^'i.i)  found  a  similar  periodic,  cyclical  variation  in  the  try- 
panosomes in  the  blood  of  sub-inoculated  rats,  guinea-pigs  and  rabbits. 
On  counting  the  parasites  in  the  blood  of  similar  animals  inoculated 
with  T.  gambiense,  they  established,  by  enumerative  methods,  that 
T.  rhodesiense  was  more  virulent  than  T.  gambiense,  while  Yorke  also 
showed  this  marked  virulence  of  T.  rhodesiense  in  practically  all 
laboratory  animals.  In  other  words  the  duration  of  infection  in  the 
case  of  T.  rhodesiense  was  shorter.  It  w^as  also  found  that  T.  rhodes- 
iense was  resistant  to  atoxyl.  The  patient,  from  whom  the  original 
strain  was  obtained,  died  about  nine  months  after  the  probable  date 
of  infection.  Some  patients  infected  with  T.  rhodesiense  have  died  in 
an  even  shorter  period,  such  as  four  or  five  months. 

In  sheep  and  goats  T.  rhodesiense  causes  an  acute  disease,  marked 
by  high  fever,  oedema  of  the  face,  and  keratitis,  as  shown  by  Bevan 
and  others,  death  resulting  after  a  relatively  short  period.  T.  gam- 
biense gives  rise,  in  these  animals,  to  no  symptoms  except  fever,  which 
may  be  overlooked.     T.  rhodesiense  produces  keratitis  in  dogs. 

Stannus  and  Yorke  (1911)  observed  T.  rhodesiense  in  animals 
inoculated  from  a  case  of  sleeping  sickness  in  Nyasaland.  Sir  D.  Bruce 
and   his   colleagues^    have    shown    (19 12)    that    T.  rhodesiense  is  the 

'  Proc.  Roy.  Soc,  B,  Ixxxii,  p.  411.  -  Annals  Trop.  Med.  and  Pm'asitol.^  iv,  p.  417, 

^  Proc.  Roy.  Soc,  B,  Ixxxv,  p.  423. 


TRYPANOSOMA    RHODESIENSE 


79 


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8o  THE   ANIMAL   PARASITES   OF   MAN 

parasite  usually  found  in  man  and  in  animals  sub-inoculated  from 
cases  of  sleeping  sickness  in  Nyasaland.  It  has  since  been  found  in 
German  East  Africa  and  Portuguese  East  Africa,  while  Ellacombe 
has  described  a  case  from  North-western  Rhodesia. 

(3)  Serum  Reactions. — Interesting  experiments  on  this  subject  were 
performed   during    1911   and  1912  by  various    French    investigators. 

{a)  Action  of  Immune  Serum  (Mesnil  and  Ringenbach)^ :  (i)  A  goat 
was  infected  with  T.  rhodesiense.  Twenty-two  days  later  its  serum 
mixed  with  T.  rhodesiense  was  injected  into  a  mouse.  Result:  Pro- 
tection. (2)  The  serum  mixed  with  T.  gamhiense  was  injected  into 
a  mouse.     Result :  Infection. 

(b)  Action  of  Baboon  Serum. — Contrary  to  T.  gambiense,  T.  rhode- 
siense is  very  susceptible  to  human  and  baboon  sera.  Mesnil  and 
Ringenbach^  showed  that  a  dose  of  i  c.c.  of  baboon  (Papio  anubis) 
serum  cured  mice  infected  with  T.  rhodesiense.  In  the  same  dose  it 
acted  very  feebly  on  T.  gambiense. 

(c)  Action  of  Human  Serum. — i  c.c.  of  human  serum  cured  T. 
rhodesiense  mice  in  three  out  of  four  cases  ;  on  T.  gambiense  mice 
there  was  no  appreciable  effect. 

Laveran  and  Nattan-Larrier^  have  shown  the  same,  namely,  that 
human  sera  act  on  T.  rhodesiense,  but  are  quite  without  action  on 
J.  gambiense. 

(d)  Trypanolytic  Reactions. — Mesnil  and  Ringenbach*  have  also 
shown  that  the  sera  of  animals  (man,  monkey  and  guinea-pig) 
infected  with  T.  gambiense  are  trypanolytic  for  the  homologous 
trypanosome,  that  is,  T.  gambiense,  but  have  no  action  on  the  hetero- 
logous trypanosome,  that  is,  T.  rhodesiense. 

(4)  Cross  Immunity  Experiments. — (a)  Mesnil  and  Ringenbach^ 
immunized  a  monkey  (Macacus  rhesus)  against  T.  gambiense.  It  was 
inoculated  with  T.  rhodesiense  on  June  7,  191 1  ;  on  June  27  trypano- 
somes  appeared,  the  infection  being  slight ;  on  July  4  it  died.  A 
control  died  in  ten  and  a  half  days. 

(b)  Laveran^  immunized  a  goat  and  mice  against  T.  gambiense. 
When  they  had  acquired  a  solid  immunity,  they  were  inoculated  with 
T.  rhodesiense.     They  became  infected  like  the  controls. 

(c)  Laveran  and  Nattan-Larrier^  immunized  a  ram  against  T.  brucei, 
it  subsequently  became  infected  with  T.  rhodesiense. 

{d)  Laveran^  immunized  a  ram  and  a  sheep  against  different  strains 
of  T.  brucei.  Inoculated  with  T.  rhodesiense  they  both  acquired  acute 
infections  and  died.     Conclusion  :  T,  rhodesiense  is  not  T.  brucei. 

'  C.R.  Soc.BioL,  Ixxii,  p.  58.  ^  (j  /^   Acad.  Sc/.,  153,  p.  1,097. 

8  C./^.  Acad.  Sci.,  154,  p.  18.  4  c.J^.  Soc.  Biol.,  Ixxi,  p.  609. 

*  C.R.  Soc.  Biol.,  Ixxi,  p.  271.  «  Bull.  Soc.  Path.  Exot.,  v,  pp.  26,  241. 

^  C.R.  Acad.  Sci.,  154,  p.  18.  «  Bull.  Soc.  Path.  Exot.,  v,  p.  loi. 


TRYPANOSOMA    KHODESIENSE  8 1 

When  the  converse  set  of  experiments  is  tried,  namely,  immuniz- 
ing an  animal  against  T.  rhodesiense,  and  then  inoculating  with 
T.  gambiense,  the  difficulty  immediately  arises  that  it  is  impossible  to 
immunize  an  animal  against  T.  rhodesiense,  owing  to  its  virulence. 
But  a  partial  and  transitory  immunity  to  T.  rhodesiense  can  be  obtained 
by  treating  the  infected  animal  with  drugs,  such  as  arsenophenylglycin. 
The  results,  so  far  as  Ihey  go,  seem  to  show  that  an  animal  immunized 
against  T.  rhodesiense  is  immune  not  only  to  T.  rhodesiense,  but  also 
to  T.  ganibiense,  a  fact  which,  according  to  Mesnil  and  Leger,  does 
not  invalidate  the  specificity  of  T.  rhodesiense,  but  tends  to  show  that 
the  two  trypanosomes  are  closely  related. 

(5)  Mode  of  Transmission  and  Reservoir. — Kinghorn  has  shown  that 
T.  rhodesiense  is  transmitted  by  Glossina  niorsitans  in  which  it  under- 
goes development.  Kinghorn  and  Yorke^  found  that  about  16  per 
cent,  of  the  wild  game  examined  in  Northern  Rhodesia  was  naturally 
infected  with  T.  rhodesiense.  The  wild  game  examined  included  water- 
buck,  hartebeest,  mpala,  bushbuck  and  warthogs.  One  native  dog  near 
the  Nyasaland  border  was  found  infected,  but  not  domestic  stock. 
Taute  doubts  whether  T.  rhodesiense  really  occurs  in  wild  game. 
Approximately  3*5  per  cent,  of  the  tsetse  flies  fed  on  infected  animals 
may  become  permanently  infected  with  T.  rhodesiense,  and  capable 
of  infecting  clean  animals.  Furthermore,  a  tsetse  fly  when  once 
infective  probably  remains  infective  for  the  rest  of   its  life. 

Kinghorn  and  Yorke,  however,  have  shown  that  climatic  condi- 
tions, namely,  those  of  temperature,  also  affect  the  infectivity  of  the 
tsetse  fly,  as  the  ratio  of  flies  capable  of  transmitting  T.  rhodesiense  to 
those  incapable  of  transmitting  the  virus  is  i  :  $34  ^^^  hot  valley 
districts  (e.g.,  Nawalia,  Luangwa  Valley,  temperature  75°  to  85°  F.), 
while  on  elevated  plateaux  (e.g.,  Ngoa,  on  the  Congo-Zambesi  water- 
shed, temperature  60°  to  70°  F.)  the  ratio  falls  to  i  :  1312. 

Mechanical  transmission  by  the  tsetse  fly  does  not  occur,  if 
a  period  of  twenty-four  hours  has  elapsed  since  the  infecting  meal. 

Developniental  Cycle  in  the  Fly. — The  period  which  elapses  between 
the  infecting  feed  of  the  flies  and  the  date  on  which  they  become 
infective  varies  from  eleven  to  twenty-five  days  in  the  Luangwa 
Valley,  according  to  Kinghorn  and  Yorke.  Attempts  carried  out  at 
laboratory  temperature  on  the  Congo-Zambesi  plateau,  during  the 
cold  season,  to  transmit  T.  rhodesiense  by  means  of  G.  morsitans 
were  always  unsuccessful.  The  developmental  cycle  of  the  trypano- 
some  in  the  fly  is  influenced  by  the  temperature  to  which  the  flies 
are  subjected  (as  stated  above).  The  first  portion  of  the  develop- 
mental cycle  proceeds  at  the  lower  temperatures  (60°  to  70°  F.),  but 
higher  temperatures  are  necessary  for  the  completion  of  the  develop- 

'  Annals  Trop.  Med.  and  Parasitol.,  vii,  p.  183. 


82         _  THE   ANIMAL   PARASITES   OF   MAN 

ment  of  the  trypanosome.  Kinghorn  and  Yorke  found  that  the 
trypanosomes  may  persist  in  the  fly,  at  an  incomplete  stage  of  their 
development,  for  at  least  sixty  days  when  the  climatic  conditions 
were  unfavourable. 

The  first  portion  of  the  developmental  cycle  of  the  trypanosome 
takes  place  in  the  gut  of  the  fly.  Invasion  of  the  saHvary  glands  of 
the  tsetse  is  secondary  to  that  of  the  intestine,  but  is  necessary  for  the 
infectivity  of  the  fly.  A  relatively  high  mean  temperature,  75°  to 
85"  F.,  is  essential  for  the  passage  of  the  trypanosomes  into  the 
salivary  glands  and  the  completion  of  their  development  therein. 

Kinghorn  and  Yorke^  state  that  the  predominant  type  of  trypano- 
some in  the  intestine  of  infected  G.  morsitans  was  a  large  broad  form, 
quite  different  from  that  which  is  most  common  in  the  salivary 
glands.  The  trypanosome  in  the  glands  resembles  the  short  form 
seen  in  the  blood  of  the  vertebrate  host.  The  authors  quoted  state 
that  both  the  intestinal  and  salivary  gland  forms  of  infective  G. 
morsitans  are  virulent  when  inoculated  into  healthy  animals. 

Bruce  and  colleagues^  have  quite  recently  (June,  1914)  published 
an  account  of  their  investigations  of  T.  rhodesiense  in  G.  morsitans 
in  Nyasaland.  (Incidentally  it  may  be  remarked  that  Bruce  considers 
T.  rhodesiense  to  be  identical  with  a  polymorphic  strain  of  T.  hnicei — 
see  pp.  83,  94).  The  development  of  T.  rhodesiense  takes  place  in  the 
alimentary  canal  and  salivary  glands,  not  in  the  proboscis,  of  the 
tsetse  fly.  In  feeding  experiments  with  laboratory  bred  flies,  as 
well  as  with  a  few  wdld  flies,  fed  on  infected  dogs  or  monkeys,  only 
8  per  cent,  of  the  flies  were  found  to  be  infected  on  dissection.  Of 
such  infected  flies,  however,  only  some  allow  of  the  complete  develop- 
ment of  the  trypanosomes  within  them,  in  other  words  only  about 
I  per  cent  of  the  flies  become  infective.  The  length  of  time  which 
elapses  before  a  fly  becomes  infective  varies  from  fourteen  to  thirty- 
one  days,  averaging  twenty-three  days,  when  kept  at  84°  F.  (29°  C). 
The  dominant  intestinal  type  of  flagellate  in  the  fly  is  that  seen  in 
the  proventriculus,  which  contains  many  long,  slender  trypanosomes. 
These  proventricular  forms  find  their  way  to  the  salivary  glands, 
wherein  crithidial  and  encysted  forms  are  seen.  They  change  into 
"blood  forms,"  which  are  short,  stumpy  trypanosomes  and  are 
infective.  ''The  infective  type  of  trypanosome  in  the  salivary  glands 
— corresponding  to  the  final  stage  of  the  cycle  of  development — is 
similar  to  the  short  and  stumpy  form  found  in  the  blood  of  the  verte- 
brate host."  The  cycle  is  thus  very  similar  to  that  of  T,  ganibiense  in 
G.  palpalis  (fig.  30). 

Culture. — ].  G.   Thomson   (1912),^  and  subsequently  Thomson 

'  Annals  Trap.  Med.  and  Parasitol.,  vii,  p.  281.         ^  Froc.  Koy.  Soc,  B,  Ixxxvii,  p.  516. 
^Annals  Trop.  Med.  and  Parasitol.,  vi,  pp.  103,  331. 


TRYPANOSOMA   CRUZI  83 

and  Sinton,  succeeded  in  cultivating  T.  rhodesiense  in  a  modified 
Novy-MacNeal  medium.  The  development  obtained  resembled  that 
of  the  trypanosome  in  the  intestine  of  Glossina, 

General  Note  on  Trypanosomes  with  Posterior  Nuclei. 

Posteriorly  placed  nuclei  have  been  found  to  occur  not  only  in 
T.  rhodesiense  by  Stephens  and  Fantham  (1910),  but  also  in  T.  pecaiidi 
by  Wenyon  {191 2),  in  T.  brucel  by  Blacklock  (191 2),  and  in  T.  eqiii- 
perdiun  by  Yorke  and  Blacklock  (191 2). 

Recently  Stephens  and  Blacklock  (1913)^  have  shown  that  two 
trypanosomeS;  different  morphologically,  have  been  confused  under 
the  name  T.  hriicei.  One  of  these  is  polymorphic  {i.e.,  it  exhibits 
long  and  slender  as  well  as  short  and  stumpy  forms)  and  came  from 
Uganda,  while  the  other  is  monomorphic  and  is  the  original  Zululand 
strain  described  by  Bruce  from  cattle  suffering  from  ^^nagana."  Bruce 
(1914)  considers  that  morphological  change  has  occurred  in  T.  brucei 
in  its  passage  through  laboratory  animals,  and  thus  explains  the 
diversity  of  views.  The  posterior  nuclear  forms  described  by 
Blacklock  occurred  in  the  Uganda  strain  of  T.  brucei.  (See  p.  95.) 
Similarly,  a  posterior  nuclear  form,  T.  equi,  has  been  separated  from 
T.  eqniperdum.     (See  p.  98.) 

Again,  Bruce  and  his  colleagues  on  the  Royal  Society  Commis- 
sion investigating  sleeping  sickness  in  Nyasaland,  have  stated  (April, 
1913)  that  ^'evidence  is  accumulating  ih-^iZ,  rhodesiense  and  T.  brucei 
(Plimmer  and  Bradford)  are  identical."  The  exact  identity  of  trypano- 
somes showing  posterior  nuclei  is,  then,  far  from  settled,  although 
Laveran  by  cross  immunity  tests  has  declared  that  T.  brucei  is  distinct 
from  T.  rhodesiense.  No  one  has  yet  seen  posterior  nuclei  in  T.gambiense. 

Trypanosonna  cruzi,  Chagas,  1909. 

Syn.  :    Schizotrvpanutn   cruzi,    Chagas,    1909. 

The  trypanosome  was  discovered  by  Chagas^  in  the  intestine  of 
the  bug,  Triatoma  (Conorhinus)  megista,  in  Brazil,  and  then  in  the 
blood  of  a  small  monkey  bitten  by  the  bug.  A  little  later  it  was  found 
in  the  blood  of  a  child,  aged  two  years,  suffering  from  irregular  fever, 
extreme  anaemia  and  enlarged  glands  in  the  State  of  Minas  Geraes, 
Brazil.  Chagas  found  that  he  was  able  to  infect  many  of  the  usual 
laboratory  animals  with  the  trypanosome,  by  allowing  the  bug  to  bite 
them.     He  was  also  able  to  culture  the  parasite  on  blood  agar. 

Chagas  found  the  Reduviid  bug,  Triatoma  megista,  in  the  houses 
of  the  poorer  inhabitants  of  the  Brazilian  mining  State,  and  that  it 
attacked  the  people,  more  especially  the  children,  at  night,  biting  the 
face.      On    this   account    the     insect    is    called    *'  barbeiro "    by   the 

'  Froc.  Roy.  Soc,  B,  Ixxxvi,  p.  187.  ^  Me7?i.  Inst.  Oswaldo  Crtiz.,  i,  p.  159. 


84 


THE   ANIMAL   PARASITES   OF   MAN 


inhabitants.  The  bite  is  somewhat  painful.  The  disease  has  since 
been  found  in  other  parts  of  Brazil,  e.g.,  Matta  de  Sao  Joao  in  Bahia 
province,  Goyaz,  Matto  Grosso  and  Sao  Paulo  provinces,  as  well  as 
in  Minas  Geraes. 

Morphology. — The  trypanosome  has  a  large  blepharoplast  or  kinetic 
nucleus.  It  is  stated  to  occur  both  free  and  in  the  red  blood  corpuscles 
in  the  peripheral  blood.     It  is  about  20  /a  long,  on  an  average. 

Two  forms  of  the  parasite  (fig.  33,  6,  7)  are  described  in  the  human 
blood.  In  one  free  form  there  is  a  large  egg-shaped  blepharoplast 
and  the  posterior  (aflagellar)  end  of  the  parasite  is  drawn  out.  The 
blepharoplast  (kinetic  nucleus)  may  have  a  chromatin  appendage. 
The  nucleus  is  oval  or  band-like,  containing  a  karyosome.  The 
flagellum,  starting  close  to  the  blepharoplast  or  its  appendage,  has  a 
free  portion  of  variable  length.  The  other  free  form  in  the  blood  has 
a  more  or  less  round,  terminal  blepharoplast,  smaller  than  in  the  first 
form,  without  a  chromatin  appendage  as  a  rule.  The  body  of  this 
second  form  is  decidedly  broader  than  that  of  the  first  mentioned. 


Fig.  33. — Jrypavosoma  cruzi.  Schizogony,  i,  merozoite  in  red  blood  corpuscle  ;  -?, 
parasite  totally  enclosed  in  red  cell,  no  flagellum  or  undulating  membrane;  j-5,  parasitts 
partially  enclosed  in  red  cell ;  6,  7,  parasites  in  human  blood  ;  8-11,  parasites  in  lungs  of  the 
monkey,  Callithrix ;  12,  jj,  initial  forms  of  schizogony ;  i^,  75,  schizogony  in  the  lungs  of 
Callithrix.     (After  Chagas.) 

The  dimorphism  has  been  interpreted  sexually,  the  first  mentioned 
forms  being  termed  males,  the  second  ones  females.  The  correctness 
of  this  interpretation  is  very  doubtful. 

No  sign  of  longitudinal  division  was  ever  seen  in  the  peripheral 
blood  or  in  the  internal  organs.  The  ''  endocorpuscular  "  forms  may 
be  completely  or  partially  enclosed  in  the  red  cell  or  only  attached 
thereto  (fig.  33,  i-S)'  ^^  the  beginning  of  infection  the  endocor- 
puscular forms  are  the  more  numerous.  Some  authorities,  however, 
doubt  these  stages. 

Life-history  in  the  Vertebrate  Host. — Chagas  found  fluctuations  in 
the  number  of  the  parasites  in  the  peripheral  blood.  He  believes  the 
increase  of  the  parasites  to  be  periodic. 


TRYPANOSOMA   CRUZI  85 

The  investigations  of  Chagas  and  of  Hartmann  have  revealed  two 
types  of  miiltiphcation  which  take  place  in  the  internal  organs  of  the 
vertebrate  host. 

(a)  The  first  type — which  possibly  belongs  to  another  organism, 
Pneiujiocystis  carinil,  see  p.  90 — occurs  in  the  capillaries  of  the  lungs. 
Tlie  flagellate  parasite  entering  the  lung  capillaries  loses  its  flagellum 
and  undulating  membrane.  Its  body  becomes  curved,  and  the  two 
ends  fuse,  and  so  an  oval  mass  is  formed  (fig.  33,  8-1 1).  In  some 
cases  the  blepharoplast  disappears,  in  other  cases  it  blends  or  fuses 
with  the  nucleus.  The  nucleus  of  the  rounded  parasite  then  divides 
into  eight  by  successive  divisions  (fig.  33,  12-15).  Next  the  body, 
which  is  surrounded  by  its  own  periplast,  also  divides,  giving  rise 
to  eight  tiny  daughter  individuals  or  merozoites  (fig.  33,  75).  The 
merozoites  lie  inside  the  periplast,  which  acts  as  a  sort  of  "  cyst 
wall."  The  merozoites  are  said  to  exhibit  dimorphism,  and  Chagas 
has  interpreted  the  dimorphism  in  terms  of  sex.  The  daughter  forms, 
produced  by  the  parent  trypanosomes  which  kept  their  blepharoplasts, 
themselves  have  blepharoplasts  as  well  as  nuclei,  and  have  been  termed 
^'males''  or  '' microgametes."  The  merozoites,  arising  from  parent 
trypanosomes  which  lost  their  blepharoplasts,  have  themselves  only 
nuclei,  and  have  been  called  ^'females"  or  ^^  macrogametes."  In  the 
case  of  the  so-called  ''female"  forms  the  single  nucleus  divides  into 
two  unequal  parts,  of  which  the  smaller  becomes  the  blepharoplast, 
and  a  flagellum  is  formed  later.  The  so-called  ''  males  "  possess  early 
a  rudiment  of  a  flagellum.  Both  kinds  of  merozoites  escape  from  the 
parent  periplast  wall,  and  enter  red  blood  corpuscles.  They  grow  into 
flagellates  within  the  corpuscles,  and  then  become 
free  as  adult  trypanosomes  in  the  blood-stream. 

(6)  The  second  mode  of  multiplication 
is  one  of  asexual  reproduction  (schizogony 
or  agamogony).  It  was  first  described  by 
Hartmann  from  hypertfophied  endothelial  cells 
of  the  lungs.  It  has  since  been  found  in  the 
cardiac  muscle,  in  the  neuroglia  of  the  central 
nervous  system,  and  in  striped  muscle  (fig.  34).  fig.  34.  —  Trypanosoma 
In  laboratory  animals  it  has  also  been  found     ^':^^''-     Transverse  section 

.....  -  .  ,  ^  of  a   striated    muscle    con- 

m    the    testicle   and  suprarenal   capsules.      In     taining  rounded  forms  of 
these  tissues  the  parasite  is  intracellular,   ap-     the  parasite  in  the  central 

,,  1,1,  •  ,  ,  portion.      X    1,000    approx. 

pearmg  as  a  small  rounded  body  with  nucleus     (After  Vianna.) 
and  blepharoplast,  without  flagellum  or  undu- 
lating membrane.     In  other  words  the  parasite  is  Leislunania-Vike  in 
the  body  tissues,  and  recalls  the  organism  of  kala-azar. 

Chagas  considers  this  second  mode  of  multiplication  to  be  strictly 
asexual.     By  this  means  the  number    of  parasites  in  the  vertebrate 


86  THE   ANIMAL   PARASITES   OF   MAN 

host  is  increased,  and  symptoms  are  produced.  On  the  other  hand 
the  first  mode  of  multiphcation,  seen  in  the  hing  capillaries,  is 
considered  by  Chagas  to  be  a  process  of  gametogony,  in  which  sexual 
forms  are  differentiated.  He  finds  that  (i)  the  adult  trypanosomes 
exhibit  a  dimorphism  in  human  blood  rarely  seen  in  artificially 
infected  guinea-pigs.  In  these  guinea-pigs  (infected  from  guinea-pigs) 
the  so-called  gametogony  in  the  lungs  is  seldom  seen.  (2)  The 
intermediate  host,  Triatoma  (Coiiorliiiitis),  becomes  infective  if  fed 
directly  on  infected  human  blood,  but  very  rarely  so  if  fed  on  guinea- 
pigs.  Chagas  is  led  to  believe  that  the  occurrence  of  sexual  forms 
constantly  in  the  blood  of  man  implies  a  greater  resistance  to  infection 
on  the  part  of  man  than  on  the  part  of  guinea-pigs  or  other  animals, 
assuming  the  general  hypothesis  that  the  formation  of  gametes 
represents  a  reaction  of  the  Protozoon  to  unfavourable  conditions. 
In  human  infection  the  number  of  parasites  is  always  less  than  in 
laboratory  animals,  and  their  presence  in  the  blood  is  transitory, 
lasting  from  fifteen  to  thirty  days  in  acute  cases.  In  many  cases 
examination  of  the  tissues  at  death  has  shown  the  presence  of  parasites 
in  patients  who  did  not  exhibit  them  in  the  general  circulation. 


Fig.  35.  —  Trypanosoma  crtizi.  Development  in  Triatoma  inegista.  i'6,  forms  found  in 
the  mid  gut  of  Triatoma  ;  7  flagellate  forms  found  in  ihe  posterior  part  of  the  gut  of  Triatoma. 
(After  Chagas.) 

Life  History  in  the  Invertebrate  Host. — About  six  hours  after  the 
ingestion  of  infected  blood  by  the  bug  {Triatoma  inegista),  the  kinetic 
nucleus  of  the  trypanosome  moves  toward?  the  nucleus,  and  the 
flagellum  is  usually  lost' (fig.  35,  7-5).  The  parasite  becomes  rounded 
and  Leishmania-Vike  (fig.  35,  j-5),  and  multiplies  rapidly  by  division. 
After  a  time,  multiplication  having  ceased,  the  rounded  forms  become 
pear-shaped  and  develop  a  flagellum  at  the  more  pointed  end. 
Crithidial  forms  (fig.  35,  7)  are  thus  produced  and  pass  into  the  intes- 
tine, where  they  multiply  and  may  be  seen  in  about  twenty-five  hours 
after  the  ingestion  of  blood.  The  crithidial  forms  may  also  be 
found  in  the  rectum  and  faeces.  The  last  stage  in  the  invertebrate 
is  a  small,  trypanosome-Iike  type,  long  and  thin  with  a  band-like 
nucleus  and  conspicuous  kinetic  nucleus.  These  parasites  are  found 
in  the  hind  gut  and  in  the  body  cavity.  They  find  their  way  into 
the  salivary  glands,  and  are  the  forms  (fig.  36)  which  are  transmissible 


TRYPANOSOMA   CRUZI  87 

to  a  new  vertebrate  host.     The  development  in  the  bug  takes  about 
eight  days  altogether,  after  which  time  the  bugs  are  infective. 

There  are  thus  three  prin- 
cipal phases   in  the    develop-  ^^^x       / 
ment  of  T.  criizi  in   Triatoma  (If       iP        ^ 
megista  :    (i)  A  multiplicative  yS^^^..^----.,^^^^^ 
phase  {Leishmania-Vike)  in  the         x_.>'^''* 

stomach     of     the     bug,     (2)     a         .    YiG.zd— Trypanosoma  cmzi.     Forms  found 
.,,.,.,,  ....         ,  in    the    salivary   glands    of    Triatoma    megista. 

cnthidial  phase,  which  is  also       (^fter  Chagas.) 

multiplicative,  in  the  hind-gut, 

and  (3)  a  trypanosome  phase,  which  is  '^  propagative,"  and  apparently 

passes  through  the  wall  of  the  alimentary  canal  into  the  body  cavity 

and  so  into  the  salivary  glands. 

Brumpt  found  that  7.  cnizi  could  live  in  Ciniex  lectnlarius,  C.  boueti, 
and  Ornithodorus  moiihata.  The  Cimex  faeces  may  be  infective. 
Blacklock  found  multiplication   of  the  parasite  in   C.  ledularius. 

Culture. — The  trypanosome  can  be  cultivated  on  Novy-MacNeal's 
blood  agar,  and  the  cultural  forms  resemble  those  described  in 
the  bug. 

Possible  Reservoir. — Chagas  thinks  that  probably  the  armadillo  or 
"  tatu  "  (Dasyptis  novemcinctus)  may  be  the  reservoir  of  T.  cruzi.  He 
also  thinks  that  Triatoma  geniadata  is  a  transmitter  ;  it  lives  in  the 
burrows  of  the  armadillo.  Other  carriers  may  be  Triatoma  infestans 
and  T.  sordid  a. 

Clinical  Features. — The  trypanosomiasis  of  Brazil,  produced  by 
T.  cruzi  and  spread  by  Triatoma  spp.  has  received  various  names, 
such  as  oppilagao,  canguary,  parasitic  thyroiditis,  and  coreotrypanosis. 
It  is  also  known  as  the  human  trypanosomiasis  of  Brazil,  South 
American  trypanosomiasis,  and  Chagas'  disease. 

Chagas^  reports  two  principal  forms — acute  and  chronic.  The 
acute  infection  is  rare,  and  is  characterized  by  increase  in  the  volume 
of  the  thyroid  gland,  pyrexia,  a  sensation  of  crackling  in  the  skin, 
enlarged  lymphatic  glands  in  the  neck,  axilla,  etc.,  while  the  liver  and 
spleen  are  increased  in  volume.  Sclerosis  of  the  thyroid  gland  is 
found  at  autopsy  and  fatty  degeneration  of  the  liver.  During  an 
attack  of  fever,  trypanosomes  are  found  in  the  blood.  The  acute 
form  was  only  observed  in  children. 

In  the  chronic  form  Chagas  reports  several  varieties  :  (a)  A 
pseudo-myxcedematous  form,  occurring  in  most  cases,  especially  up 
to  the  age  of  15.  There  is  hypertrophy  of  the  thyroid  gland  or 
at  least   signs   of    hypothyroidism,    general    hypertrophy   of    glands, 

^  Brazil  Medico,  Nov.  15,  1910.  Longer  account  in  Mem.  Inst.  Oswaldo  Cruz,  iii, 
pp.  219-275.     See  Sleep.  Sick.  Bull.y   Nos.   35   and  40. 


88  THE   ANIMAL   PARASITES   OF   MAN 

disturbance  of  heart  rhythm,  and  nervous  symptoms.  (6)  The  myxoe- 
dematous  form  is  characterized  by  similar  symptoms,  especially  by 
considerable  swelling  of  the  thyroid  body,  and  myxoedema  of  the 
subcutaneous  cellular  tissue  ;  sometimes  there  is  a  true  pachydermic 
cachexia,  (c)  In  the  nervous  form  there  are  motor  disturbances, 
aphasia,  disturbances  of  intelligence  or  signs  of  infantilism,  athetosis 
of  the  extremities  and  idiocy.  There  are  also  paralytic  symptoms  of 
bulbar  origin,  disturbances  of  mastication,  phonation  and  deglutition, 
and  in  some  cases  convulsive  attacks,  (d)  The  cardiac  form,  charac- 
terized by  disturbance  of  the  heart  rhythm.  In  all  these  forms  the 
parasite  is  found  at  autopsy  in  the  nervous  substance,  brain,  bulb  and 
heart. 

Vianna  (1911)^  has  studied  the  histopathology  of  the  disease. 
Some  of  the  chief  points  are  :  in  the  heart  muscle  destruction  of 
the  sarcoplasm,  followed  by  interstitial  myocarditis;  in  the  central 
nervous  system  invasion  of  the  neuroglia  cells  and  inflammatory 
reaction  ;  in  the  suprarenal  capsule  invasion  of  medulla  or  cortex  ; 
inflammatory  reaction  can  also  be  seen  in  the  kidneys,  the  hypophysis 
and  thyroid  gland. 

Recently  Chagas  states"  that  "  schizotrypanosomiasis "  has  been 
found  in  a  child  15  to  20  days  old,  and  that  Trypanosoma  criizl 
has  also  been  found  in  a  foetus — the  mother  being  infected  with 
the  trypanosome.  The  trypanosomiasis  can,  then,  be  transmitted 
hereditarily. 

Trypanosoma  lewisi,  Kent,  1881. 

The  trypanosome  has  a  nucleus  somewhat  displaced  anteriorly,  about 
one-third  of  the  way  from  the  anterior  (flagellar)  end  of  the  body,  a 
relatively  straight  edge  to  the  undulating  membrane,  and  a  rod-shaped 
blepharoplast  (fig.  37,  A).    It  averages  about  25  jju  long  and  i'5  fju  broad. 

Much  attention  has  been  devoted  in  recent  years  to  the  elucida- 
tion of  the  life  history  of  the  rat  parasite,  Trypanosoma  lewisi.  It  is 
usually  non-pathogenic  to  its  host.  It  has  been  shown  that  the 
trypanosome  can  be  transmitted  from  rat  to  rat  by  the  rat-flea, 
Ceratophyllns  fasciatus,  and  by  Ctenocephalus  canis  (the  so-called  dog- 
flea).  (See  also  p.  92).  The  flagellate  may  also  persist,  but  doubtfully 
develop,  in  the  rat-louse,  Ha^matopinns  spinnlosns.  These  researches 
may  now  be  summarized. 

Life  Cycle  in  the  Vertebrate  Host. — After  infection  of  a  rat,  the 
trypanosomes  usually  appear  in  the  animal's  blood  in  five  to  seven 
days.  This  incubation  period  applies  either  to  a  natural  or  an  artificial 
infection.  The  trypanosomes  first  observed  in  the  rat's  blood  are 
diverse  in  form  (fig.  37),  being  small,  medium  and  large  in  size.  This 
diversity  is  explained  by  the   rapid  multiplication    taking  place.      A 

'  Mem.  Inst.  Oswaldo  Cruz,  iii,  p.  276.  ^  ^^^.  Med.  S.  Paulo  (1912),  xv,  p.  337. 


TRYPANOSOMA    LEWISI 


89 


trypanosome  may  divide  by  equal  longitudinal  fission  (fig.  37,  C,  D), 
but  more  commonly  multiple  fission  occurs  (fig.  37,  G,  H),  and  is 
unequal.  Rosette  forms  are  produced,  in  which  the  parent  form  can 
be  recognized  by  its  long  flagellum  (fig.  37,  H)  and  attached  to  it  are 


Fig.  37. — Trypanosoma  lewisi,  from  rat's  blood.  A,  ordinary  form;  B,  small  form; 
C,  D,  stages  in  equal  binary  fission  ;  e,  elongate  form  {longocaudense  type),  resulting  from 
division  as  seen  in  d  ;  f,  unequal  binary  fission  ;  G,  H,  multiple  fission  into  four  and  eight  ; 
I,  small  form  ;  j,  binary  fission  of  small  form ;  K,  division  rosette.  X  2,coo.  (After 
Minchin  and   Thomson.) 

daughter  individuals,  smaller  in  size,  from  which  flagella  are  growing. 
Minchin  and  |.  D.  Thomson  (1912)  find  that  the  daughter  forms  may 
be  set  free  sometimes  with  a  crithidia-like  facies  (fig.  37,  I),  the 
blepharoplast  being  anterior  but  near  to  the  nucleus.     The  daughter 


90  THE   ANIMAL   PARASITES   OF   MAN 

forms,  when  set  free,  may  themselves  divide  by  binary  or  muUiple 
fission,  in  the  latter  case  forming  rosettes  (fig.  37,  K).  Rosette 
forms  were  described  by  Moore,  Breinl  and  H indie  in  1908. 

Lingard,  some  years  ago,  described  as  a  distinct  species,  T.  longo- 
caudense,  certain  forms  wnth  markedly  elongate  posterior  ends 
(fig.  37,  E).  According  to  Minchin,  ''  these  forms  appear  to  arise  by 
binary  fission"  (fig.  37,  D).  These  long  drawn-out  forms  ''are  of 
constant  occurrence  and  very  numerous  at  a  certain  stage  of  the 
multiplication  period."  It  is  about  the  eighth  or  tenth  day  after 
infection  that  the  multiplication  of  T.  lewisi  is  at  its  maximum  in  the 
rat's  blood.  About  the  twelfth  or  thirteenth  day  the  trypanosomes 
seen  in  the  blood  appear  uniform.  According  to  Minchin  (1912)  ^ 
the  rat  ''gets  rid  of  its  infection  entirely  sooner  or  later,  without 
having  suffered,  apparently,  any  marked  inconvenience  from  it,  and 
is  then  immune  against  a  fresh  infection  with  this  species  of  trypano- 
some."  There  is,  then,  a  cycle  of  development  in  the  vertebrate  host. 
Minchin  notes  that  the  records  of  the  pathogenicity  of  T.  leivisi  in 
rats,  causing  their  death,  need  further  investigation. 

T.  lewisi  inoculated  into  dormice  {Myoxus  iiitela)  and  jerboas  may 
become  pathogenic  thereto. 

Carini  found  cysts  in  the  lungs  of  rats  infected  with  T.  lewisi. 
He  thought  the  cysts  were  schizogonic  stages  of  the  trypanosome, 
comparable  with  those  found  in  the  lungs  of  animals  sub-inoculated 
with  T.  criisi.  Delanoe  (1912)^  has  found,  however,  that  such 
cysts,  containing  eight  vermicules,  occurred  in  rats  uninfected  with 
T.  lewisi.  Delanoe  concludes  that  the  pneumocysts  are  independent 
of  r.  leivisi f  and  represent  a  new  parasite,  Pneuinocyslis  carinii.  The 
pneumocysts  may  be  allied  to  the  Coccidia,  and  must  be  considered 
when  investigating  the  life-cycle  of  a  trypanosome  in  a  vertebrate 
host.     Some  of  the  stages  of  T.  cruzi  may  possibly  be  of  this  nature. 

Life-cycle  in  the  Invertebrate  Host. — This  occurs  in  fleas,  and  has 
been  investigated  in  considerable  detail  by  Minchin  and  Thomson 
in  Ceratophyllns  fasciatns,  and  by  Noller  in  Ctenocephalns  canis  and 
Ctenopsylla  ninscnli. 

When  infected  rat's  blood  is  taken  up  by  the  flea,  the  parasites  pass 
wMth  the  ingested  blood  direct  to  the  mid-gut  of  the  Siphonapteran. 
In  the  flea's  stomach  they  multiply  in  a  somewhat  remarkable  manner, 
namely,  by  penetration  of  the  cells  of  the  lining  epithelium,  and 
division  inside  the  epithelial  cells.  Inside  these  lining  cells  the 
trypanosomes  first  grow  to  a  large  size  and  then  form  large  spherical 
bodies,  within  which  nuclear  multiplication  occurs  (fig.  38,  A — F). 
Any  one  of  these  large  spherical  bodies  contains  at  first  a  number  of 
nuclei,  blepharoplasts  and  developing   flagella,  the  original  flagellum 

*  **  Protozoa,"  p.  294.  2  c.  R.  Acad.  Sci.,  civ,  p.  658. 


TRYPANOSOMA    LEWISI 


91 


still  remaining  attached  for  a  time.  The  cytoplasm  then  divides  into 
daughter  trypanosomes  which  are  contained  within  an  envelope, 
formed  by  the  periplast  of  the  parent  parasite.  Inside  the  periplast 
envelope  are  a  number  of  daughter  trypanosomes  "  wriggling  very 
actively ;  the  envelope  becomes  more  and  more  tense,  and  finally 
bursts  with  explosive  suddenness,  setting  free  the  flagellates,  usually 
about  eight  in  number,  within  the  host-cell  "  (fig.  38,  F).  The  daughter 
forms  escaping  from  the  host  cell  into  the  stomach  lumen  of  the  flea 
are  fully  formed,  long  trypanosomes. 


Fig.  38. —  Trypanosoma  lewisi.  Developmental  stages  from  stomach  of  rat  flea.  O, 
ordinary  blood  type  ;  A — F,  stages  occurring  in  gut-epithelium  of  flea,  when  the  trypanosome 
becomes  rounded  and  undergoes  multiplication,  forming  in  F  eight  daughter  trypanosomes  ; 
G,  type  of  trypanosome  resulting  from  such  division  which  passes  back  to  the  rectum. 
X  2,000.     (After  Minchin.) 


The  trypanosomes  (fig.  38,  G)  pass  into  the  flea's  rectum.  The 
next  phase  is  a  crithidial  one.  The  parasites  become  pear-shaped, 
in  which  the  blepharoplast  (kinetic  nucleus)  has  travelled  anteriorly 
past  the  nucleus  towards  the  fiagellum  (fig.  39).  The  crithidial  forms 
attach  themselves  to  the  wall  of  the  rectum,  and  multiply  by  binary 
fission  (fig.  39,  D).  A  stock  of  parasites  is  thus  formed  which, 
according  to  Minchin  and  Thomson,  ^^  persist  for  a  long  time  in 
the  flea — probably  under  favourable  conditions,  for  the  whole  life 
of  the  insect  "  (fig.  39,  A — I). 

From  the  crithidial  forms  of  the  rectum,  according  to  Minchin, 
small  infective  trypanosomes  arise  by  modification  morphologically 
(fig.  39,  ]— M).  The  flagellum  grows  longer  and  draws  out  more  the 
anterior  part  of  the  body,  the  blepharoplast  migrates  posteriorly, 
behind  the  nucleus,  and  carries  with  it  the  flagellar  origin.  These 
trypanosomes  are  small,  but  broad  and  stumpy  (fig.  39,  N),  and  can 


92 


THE   ANIMAL   PARASITES   OF   MAN 


infect  a  rat.  Minchin  and  Thomson  formerly  considered  that  the 
small,  stumpy,  infective  trypanosomes  pass  forwards  from  the  rectum 
into  the  stomach,  and  "  appear  to  be  regurgitated  into  the  rat's  blood 
when  the  flea  feeds."  However,  the  small  infective  trypanosomes 
were  previously  described  by  Swellengrebel  and  Strickland.^  They 
may  be  found  in  the  flea's  faeces.  Noller  (191 2)^  has  found  that  the 
development  of  T.  lewisi  proceeds  quite  well  in  the  dog  flea  (Cteno- 
cephalns  canis)  in  Germany.  Wenyon  confirms  this,  and  states  that 
the  human  flea,  Piilex  irritans,  and  the  Indian  rat-flea,  Xenopsylla 
cheopis,  are  also  able  to  serve  as  true  hosts  for  T.  lewisi. 

Noller  stated  that  rats  were  not  infected  with  T.  lewisi  by  infective 
fleas  biting  them,  but  by  the  rats  licking  up  the  faeces  passed  by  the 


^  Fig.  39. —  Trypanosoma  lewisi.  Developmental  stages  from  rectum  of  rat-flea.  A,  early 
rectal  form;  C,  D,  division  of  crithidial  foim;  E,  group  of  crithidial  forms;  F — I,  crithidial 
forms  without  free  flagella,  some  becoming  rounded  ;  J — M,  transitional  forms  to  tiypanosome 
type  seen  in  N,  which  represents  the  final  foim  in  the  flea,      x  2,000.     (After  Minchin.) 


fleas  while  feeding.  This  is  not  in  agreement  with  Minchin  and 
Thomson's  earlier  views  of  regurgitation,  which,  apparently,  they 
have  now  abandoned.^  Wenyon  (1912)  confirms  Noller's  experi- 
ments. He  took  a  dog  flea,  containing  infective  trypanosomes  in 
its  faeces,  and  allowed  it  to  feed  on  a  clean  rat.  The  faeces  of  the 
flea,  passed  while  feeding,  were  carefully  *^  collected  on  a  cover  glass 
and  taken  up  in  culture  fluid  with  a  fine  glass  pipette."  The  contents 
of  the  pipette  were  discharged  into  the  mouth  of  a  second  clean  rat. 
Injury  to  the  rat's  mouth  was  carefully  avoided.  The  first  rat,  on 
which  the  infective  flea  was  fed,  did  not  become  infected,  while  the 


'  Parasitology,  iii,  p.  360.  "^  Arch.  f.  Frotistenkunde,  xxv,  p.  386. 

^  Report  to  Advis.  Comm.  Trop.  Dis.  Research  Fund  for  1913,  p.  7-^. 


TRYPANOSOMA   BRUCEI 


93 


second  rat,  in  whose  mouth  infective  flea  faeces  were  placed,  became 
infected  in  six  days. 

When  infective  forms  of  T.  lewisi  have  been  developed  within  the 
^iit  of  a  rat  flea,  they  may  enter  and  infect  the  vertebrate  host 
by  ^  {a)  being  crushed  and  eaten  by  the  rodent ;  (6)  the  rat  may  lick 
its  fur  on  which  an  infected  flea  has  just  passed  infective  excrement ; 
or  (c)  the  rat  may  lick,  and  infect  with  flea  excrement,  the  wound 
produced  by  the  bite  of  the  flea. 

The  time  taken  for  the  full  development  of  T.  lewisi  in  the  flea 
is  about  six  days.  The  intracellular  phase  is  at  its  height  about 
the  end  of  the  first  day ;  the  crithidial  phase,  in  the  flea's  rectum, 
begins  during  the  second  day;  the  stumpy,  infective  trypanosomes 
are  developed  in  the  rectum  about  the  end  of  the  fifth  day. 

Wenyon^  writes  that,  ''the  fleas,  when  once  infected  with  T.  lewisi^ 
remain  infected  for  long  periods,  for  though  many  small  infective 
trypanosomes  are  washed  out  of  the  gut  at  each  feed,  those  that 
remain  behind  multiply  to  re-establish  the  infection  of  the  hind  gut. 
Further,  the  infection  is  still  maintained  even  if  the  flea  is  nourished 
on  a  human  being,  so  that  fresh  human  blood  does  not  appear  to  be 
destructive  to  the  infective  forms  in  the  flea." 

The  best  method  of  controlling  fleas  during  experiments  is  that 
due  to  Noller.  He  adopted  the  method  of  showmen  who  exhibit 
performing  fleas,  and  secure  them  on  very  fine  silver  wire. 

Of  fleas  fed  on  an  infected  rat  only  about  20  per  cent,  become 
infective.  About  80  per  cent,  are  immune.  If  fleas  are  examined 
twenty-four  hours  after  feeding,  trypanosomes  will  be  found  in  all,  so 
that  many  of  the  parasites  are  destined  to  degenerate. 

It  may  be  of  interest  to  note  that  Gonder^  (1911)  has  shown  that 
a  strain  of  T.  lewisi  resistant  to  arsenophenylglycin  loses  its  resistance 
after  passage  through  the  rat-louse,  Hceniatopiniis  splnulosus.  These 
experiments  suggest  that  physiological  "  acquired  characters "  may 
be  lost  by  passage  through  an  invertebrate  host. 

Trypanosoma  brucel,  Plimmer  and  Bradford,  1899. 

Trypanosoma  brucel  was  discovered  by  Sir  D.  Bruce  in  1894  in 
cattle  in  Zululand  and  was  named  T.  brucel  by  Plimmer  and  Bradford 
in  1899  in  honour  of  its  discoverer.  This  trypanosome  is  of  con- 
siderable economic  importance,  as  it  is  responsible  for  the  fatal 
tsetse  fly  disease,  or  '' nagana/'  in  cattle,  horses  and  dogs.  The 
disease  is  widely  distributed  in  Africa  and  is  transmitted  from  host  to 
host  by  the  tsetse,  Glossina  morsitans,  and  other  species  of  Glosslna, 

'  Nuttall,  Parasitology,  v,  p.  275. 

"^  Report  to  Advis.  Comm.  Trop.  Dis.   Research  Fund,  October,  1912,  p.  91.     See  also 
/ourn.  Lond.  Sch.  Trop.  Med.,  ii,  p.  119. 
3  Centralbl.f.  Bakt.,  Orig.,  Ixi,  p.  102. 


94  THE   ANIMAL   PARASITES   OF   MAN 

The  virus  is  maintained  in  nature  in  certain  big  game,  such  as  wilde- 
beest, bushbuck  and  koodoo,  which  thus  act  as  Uving  reservoirs  of 
disease  from  which  the  tsetse  may  become  infected.  These  reservoir 
hosts  are  not  injured,  apparently,  by  the  presence  of  the  parasites. 

T.  hriicei  is  rapidly  fatal  to  the  small  laboratory  animals,  such  as 
rats  and  mice.  Horses,  asses  and  dogs  practically  always  succumb 
to  its  attacks,  while  a  very  small  number  of  cattle  recover  from 
'^  nagana."  The  disease  is  characterized  by  fever,  destruction  of  red 
blood  corpuscles,  severe  emaciation  and  by  an  infiltration  of  coagu- 
lated lymph  in  the  subcutaneous  tissue  of  the 
neck,  abdomen  and  extremities  giving  a  swollen 
appearance  thereto.  The  natural  reservoirs  in 
which  T.  hriicei  has  been  long  acclimatized  are 
unaffected  by  the  trypanosomes,  while  the  newer 
hosts,  such  as  imported  cattle  in  Africa,  are 
rapidly  destroyed  by  their  action. 

The  general  morphology  and  life  history  in 
the  vertebrate  host  is  that  of  a  typical  trypano- 
some  (fig.  40).  Its  length  is  from  12  /t  to  35  /a, 
its  breadth  from  1*5  /a  to  4/x.  Multiplication  by 
longitudinal  division  proceeds  in  the  peripheral 
blood  (fig.  26),  while  latent,  leishmaniform 
Fig.  ^o.— Trypanosoma  bodics  are  produced  in  the  internal  organs. 
iTv'etn  a„d'^;„il.t''"  Bruce  and  colleagues'  have  quite  recently 

(June,  1914)  described  the  development  of  a 
Zululand  strain  of  T.  hrucei  in  G.  morsitans.  The  tsetse  flies  were 
bred  out  in  Nyasaland.  In  vertebrate  blood  the  brncei  strain 
was  polymorphic.  The  development  was  like  that  found  for 
T.  gambiense  in  G.  palpalis  (fig.  30),  and  by  Bruce  and  colleagues  for 
T.  rhodesiense  in  G.  morsitans  in  Nyasaland.  Long  trypanosomes 
were  found  in  the  proventriculus  of  the  tsetse.  Crithidial,  rounded 
or  encysted,  and  immature  ^'  blood  forms "  occurred  in  the  salivary 
glands;  and  finally  infective,  stumpy,  ''blood  forms  "  were  differen- 
tiated in  the  salivary  glands.  The  period  of  development  of  T.  hrucei 
in  G.  morsitans  takes  about  three  weeks,  and  then  the  fly  becomes 
infective.  Bruce  believes  that  T.  rhodesiense  of  Nyasaland  and 
T.  hrucei  of  Zululand  are  the  same,  their  cycles  of  development 
in  G.  morsitans  being  "  marvellously  alike."  (But  see  Laveran,  p.  80.) 
T.  hrucei  has  been  cultivated  with  difficulty  by  Novy  and  MacNeal, 
using  blood  agar.  The  best  treatment  for  nagana  is  arsenic  in  some  form. 
It  is  probable  that  more  than  one  trypanosome  has  been  con- 
fused under  the  name  T.  hrucei,  more  especially  as  the  occurrence  of 
many  species  of  trypanosomes  in  various  animals  in  Africa  was  not 

'  Proc.  Roy.  Soc.^  Bj  Ixxxvii,  p.  526. 


TRYPANOSOMA   EVANSI  95 

suspected  until  comparatively  recent  times.  It  has  been  shown  by 
Stephens  and  Blacklock  (191 3)  that  the  original  Zululand  strain  of 
T.  hnicei  was  monomorphic,  while  the  organism  sent  from  Uganda, 
and  at  the  time  believed  by  Bruce  to  be  the  same  as  the  Zululand 
trypanosome,  has  been  found  to  be  polymorphic,  with  morpho- 
logical resemblances  to  T.rJiodesiense.  Stephens  and  Blacklock^  have 
suggested  the  name  T.  iigandce  for  the  polymorphic  trypanosome, 
which,  however,  has  marked  resemblances  with  Trypanosoma 
pecaudi,  and  they  are,  perhaps,  identical.  T.  pec  audi  was  the  name 
given  by  Laveran  ^  in  1907  to  the  causal  agent  of  '*  baleri "  in  equines 
and  sheep  in  the  French  Sudan.  T.  pecaudi,  which  is  dimorphic, 
is  widely  distributed  in  Africa.  An  extremely  small  number  of  both 
T.  pecaudi  and  T.  ugaudce  have  been  shown  to  possess  posterior 
nuclei.  T.  pecaudi  is  transmitted  by  various  species  of  Glossina, 
and  is  said  to  develop  in  the  gut  and  proboscis  of  the  fly. 

On  the  other  hand,  Bruce  and  colleagues  (1914),  examining  a 
strain  sent  from  Zululand  in  1913,  state  that  T.  hrucei  is  polymorphic. 
Bruce  (1914)  suggests  that  passage  through  laboratory  hosts  has 
influenced  and  altered  the  morphology  of  the  parasite. 

Trypanosoma  evansi,  Steel,  1885. 

Syn. :   Spirochceta  evansi^  Steel,    1885;   Hcematomonas  evansi,  Crookshank,   1886; 
Tf'ichomonas  evansi^  Crookshank,  1886. 

Trypanosoma  evansi,  first  found  by  Evans  in  1880,  in  India,  is  the 
causal  agent  of  the  disease  known  as  **  surra."  The  malady  affects 
more  particularly  horses,  mules,  camels  and  cattle  in  India  and  neigh- 
bouring countries,  such  as  Burma  and  Indo-China.  It  occurs  also  in 
Java,  the  Philippines,  Mauritius  and  North  Africa.  Elephants  may 
be  affected.  A  serious  outbreak  among  cattle  in  Mauritius  occurred 
in  1902,  the  disease  being  imported  into  the  island.  The  symptoms 
are  fever,  emaciation,  oedema,  great  muscular  weakness  and  paralysis 
culminating  in  death. 

T.  evansi  varies  from  18  //,  to  34  //<  in  length  and  i'5  //-  to  2  //,  in 
breadth.  It  has  a  pointed  posterior  extremity,  and,  anteriorly,  there 
is  a  free  portion  to  the  flagellum  (fig.  41).  It  is  possibly  mono- 
morphic, but  a  few  broad  forms  occur.  The  trypanosome  multiplies 
by  longitudinal  fission  in  the  blood.  Rounded  leishmaniform  stages 
occur  in  the  spleen  of  the  vertebrate  host,  which  stages  Walker^  (1912) 
considers  to  be  phases  of  schizogony. 

The  parasite  is  transmitted  in  nature  by  various  species  of  Tabanus 
and  Stomoxys,  though  at  present  little  is  known  of  the  life-history 
within  these  invertebrate  hosts. 

'  Froc.  Roy.  Soc.^  B,  Ixxxvi,  p.  187.  "^  C.R.  Acad.  Set.,  cxliv,  p.  243. 

■•*  Philippine  lourn.  Sc.  (Sect.  B),  vii,  p.  53. 


96 


THE   ANIMAL   PARASITES   OF   MAN 


Fig.  41.  —  Trypanosoma 
evaiisi.  x  2,000.  (Original. 
From  preparation  by  Fan- 
tham.) 


Dogs   are   said  to    contract   the   disease    by   feeding   on    animals 
dead  of  surra. 

A  variety  of  T.  evansl  is  the  cause  of  ''  mbori  "  in    dromedaries 

in  Africa  (Sahara  and   Sudan).     Another  possible  variety,  or  closely 

allied  form,  is  T.  soudaiiense,  the  causal  agent 

of  ''el  debab  "  in  camels  and  horses  in  North 

Africa,  especially  Algeria  and  Egypt. 

An  extraordinary  example  of  the  possible 
infection  of  a  human  being  with  an  animal 
trypanosom.e  is  recorded  in  the  case  of  Pro- 
fessor Lanfranchi,  of  the  Veterinary  School, 
Parma.  The  Professor  became  infected  with 
trypanosomes,  although  only  nagana  and  surra 
were  maintained  in  his  laboratory,  and  he 
himself  had  never  visited  the  tropics.  He 
suffered  from  irregular  attacks  of  fever  and 
was  cedematous,  but  his  mind  remained  clear. 
The  identification  of  the  trypanosome  from 
Lanfranchi's  blood  has  been  a  matter  of  great 
difficulty.  Apparently  Mesnil  and  Blanchard 
(1914)^  consider  the  strain  found  in  the  patient  is  almost  indistinguish- 
able in  its  reactions  from  T.  gainbiense,  though  the  parasite  is  mono- 
morphic.     Lanfranchi  considers  that  he  was  infected  with  T.  evansi. 

Trypanosonna  equinum,  Voges,  1901. 

Syn.  :   Trypafwsoina  ehnassiaiH^  Lignieres. 

Trypanosoma  eqiiinum  was  found  by  Elmassian  to  be  the  cause  of 
the  fatal  disease,  '' mal  de  caderas,"  of  horses 
and  dogs,  in  South  America  (Paraguay,  Argen- 
tine, Bolivia).  The  name  refers  to  the  fact  that 
in  the  disease,  as  in  other  trypanosomiases, 
the  hind  quarters  become  paralysed.  Cattle 
are  refractory  to  inoculation. 

T.  equinum  is  about  22  yu,  to  24  /x  long 
and  about  1*5  fi  broad  (fig.  42).  Although  this 
trypanosome  is  very  active,  yet  it  is  charac- 
terized by  the  blepharoplast  (kinetic  nucleus) 
being  very  minute  or  even  absent,  as  the 
granule    sometimes    seen    may    be    the    basal 

granule   of   the    fiagellum.  Fig.   42.  —  Trypanosoma 

The  mode   of  transmission  of  T.  equinum    ^qt^innm.    x  2,000.    (After 

,   ,  -ji       1        1    J  i    •     i  TVT-  Laveran  and  Mesnil.) 

is  not  known  with  absolute  certainty.     Migone 

has  shown  that  the  parasite  causes  a  fatal  disease  in  the  large  South 


*  Bull.  Soc.  Path.  Exot.,  vii,  p.  196. 


TRYPANOSOMA  EQUIPERDUM  97 

American  rodent,  the  capybara  {Hydrochcenis  capybara).  This  animal 
appears  to  be  a  reservoir  of  the  parasite.  Dogs  may  become  infected 
by  eating  diseased  capybaras,  and  it  is  suggested  that  the  infection  is 
spread  from  the  dogs  to  horses  by  the  agency  of  fleas.  Some  authorities 
consider  that  T.  equimtin  may  be  spread  by  various  Tabanidce  and  by 
Stomoxys.  Neiva  (1913)^  doubts  all  these  modes  of  transmission  in 
Brazil,  and  suggests  Chrysops  or  Trlatoma  as  vectors. 

Trypanosoma  equiperdum,  Doflein,  1901. 
Syn. :   Trypanosoma  rougeti,  Laveran  and  Mesnil. 

The  malady  of  horses  known  as  "  dourine  "  or  ''  mal  du  coit "  is 
due  to  a  trypanosome,  T,  equiperdtun,  discovered  by  Rouget  in  1894. 
"Dourine" — also  known  as  ^'stallion  disease"  or  '^covering  disease" 
— is  found  among  horses  and  asses  in  Europe, 
India,  North  Africa  and  North  America.     The 
trypanosome  is  transmitted  by  coitus,  and  so 
far  as  is  known  not  by  insect  agency. 

The  progress  of  the  disease  may  be  con- 
sidered under  three  periods.  The  period  of 
oedema,  when  signs  of  oedema  of  the  genitalia 
are  seen.  The  oedema  is  generally  painless 
and  non-inflammatory.  This  period  lasts  about 
a  month.  It  is  succeeded  by  the  period  of 
eruption,  which  sets  in  about  two  months 
after  infection.  Circular  oedematous  areas 
(*'  plaques  "),  often  about  the  size  of  a  two-  fig.  43.  —  Trypamsowa 
shilling  piece,  appear  under  the  skin  of  the  sides     equiperduvi.  x  2oooapprox- 

,  ,  .     \  '^^  1       ,,  imately.     (Original.      From 

and  hmd  quarters,  and  also,  at  times,  under  the     preparation  by  Famham.) 
skin  of  the  neck,  thighs  and  shoulders.     The 

eruption  is  variable,  but  usually  lasts  about  a  week  and  leaves  the 
animal  in  an  enfeebled  condition.  Gland  enlargement  and  swelling 
of  the  joints  and  synovia  also  may  occur.  The  third  period  of  the 
disease  is  described  as  that  of  ancemia  and  paralysis.  The  animal 
becomes  very  anaemic,  emaciation  is  marked,  superficial  non-healing 
abscesses  often  form,  and  conjunctivitis  and  ulcerative  keratitis  can 
occur.  Paralysis  ensues,  and  in  from  two  to  eighteen  months  the 
animal  dies.  In  the  acute  form  of  the  disease  the  animal  may  die 
after  the  first  period  from  acute  paralysis. 

It  is  difficult  to  find  the  trypanosomes  in  naturally  infected 
animals,  and  they  are  best  obtained  from  the  plaques  of  the 
eruption.  Apparently  the  parasite  occurs  more  in  the  lymph  than 
in  the  blood. 

'  Brazil  Medico^  xxvii,  p.  366. 


98  THE   ANIMAL   PARASITES   OF   MAN 

Ruminants  are  said  to  be  refractory  to  this  trypanosome. 

T.  equiperdum  is  about  25  yu,  to  28  /a  in  length  on  an  average,  but 
varies  from  16  /x  to  35  fjb.  Its  cytoplasm  is  relatively  clear,  and  does  not 
show  chromatic  granules  (fig.  43).     It  is  stated  to  be  monomorphic. 

It  has  been  shown  recently  by  Blacklock  and  Yorke  {1913)^  that 
there  is  another  trypanosome  giving  rise  to  dourine  in  horses.  This 
trypanosome  is  dimorphic  (resembling  T.  pecandi  and  7.  iigandce), 
and  is  named  T.  eqni.  Previously  T.  equiperdum  and  T.  eqtii  had 
been  confused. 

Uhlenhuth,  Hiibner  and  Worthe  have  demonstrated  the  presence 
of  endotoxins  in  T.  equiperdum.  These  endotoxins  may  be  set  free 
by  trypanolysis. 

Trypanosoma  theiteri,  Bruce,  1902. 

This  parasite,  60  /x  to  70  //-  long,  and  4  /i  to  5  /a  broad,  is  dis- 
tinguished for  its  large  size,  though  it  is  not  so  large  as  7.  ingens 
from  Uganda  oxen,  whose  length  may  be  72  //,  to  122  fi,  and  breadth 

7  /A  to  10  fjL.  The  posterior 
end  of  T.  theileri  is  drawn 
out.  Small  forms  of  the  flag- 
ellate are  known,  25  /x,  to  53  /x 
in  length.  Probably  other 
forms  of  the  parasite  have  the 
nucleus  posterior,  and  these 

Fig.  44.  -  Trypanosoma  theileri.      x  2,000.  fl o  rr^l  1  a f  pc  wpi-p  f  nrm pri  v  c;pn 

(After  Laveran  and  Mesnil.)  tlagcllates  wei c  toi  meriy  Sep- 

arated as  r.  transvaaliense 
(Laveran,  1902).  Myoneme  fibrils  may  be  seen  on  its  body.  The 
pathogenicity  of  this  organism  is  doubtful,  it  was  formerly  thought 
to  be  the  causal  agent  of  ^'  gall-sickness  "  in  cattle  in  South  Africa. 
T.  theileri  also  occurs  in  Togoland,  German  East  Africa,  and  Trans- 
caucasia.    Allied  or  identical  parasites  occur  in  cattle  in  India. 

Trypanosoma  theileri,  specific  to  cattle,  is  perhaps  transmitted  by 
the  fly  Hippobosca  rufipes  in  South  Africa, 

Trypanosoma  hippicum,  Darling,  1910. 

Trypanosoma  hippicum  causes  the  disease  of  mules  known  as 
'*murrina."2  It  was  found  in  mules  imported  to  Panama  from  the 
United  States.  It  can  live  in  other  equines.  The  parasite  varies  from 
18  /x  to  28  /A  in  length,  and  is  from  1*5  ytt  to  3  //.  broad.  Its  undu- 
lating membrane  is  little  folded.  The  trypanosome  has  a  noticeable 
blepharoplast.  It  can  penetrate  mucous  membranes,  and  it  is  thought 
that  the  trypanosome  may  be  transmitted  during  coitus.     It  may  also 

^  Proc.  Roy.  Soc,  B,  Ixxxvii,  p.  89.  ^  Bull.  Soc.  Path.  Exot.,  iii,  p.  381. 


TRYPANOSOMA   VIVAX  99 

be  spread  meclianically  by  species  of  Miisca,  Sarcophaga  and  Comp- 
somyia,  sucking  the  wounds  of  infected  animals  and  carrying  over  the 
trypanosomes  to  wounds  on  healthy  ones. 

Endotrypanum  schaudinni,  Mesnil  and  Brimont,  1908. 
This  organism  was  discovered  in  the  blood  of  a  sloth  {Choloepus 
didactyliis),  m  South  America  (French  Guiana).^  It  possesses  special 
interest,  in  that  the  best  known  form  of  the  organism  is  endoglobular, 
inhabiting  the  erythrocytes  of  the  sloth.  A  free  trypanosome  in  the 
same  animal  was  considered  to  be  different  from  the  endoglobular 
form,  which  w^as  somewhat  like  a  peg-top,  and  possessed  a  short 
fiagellum.  Darling^  (November,  1914)  has  seen  the  organism  in 
Panama.  He  describes  free  crithidial  forms  in  shed  blood,  but  not 
in  the  blood-stream  of  the  sloth. 

Trypanosoma  boylei,  Lafont,  1912. 

This  is  a  parasite  of  the  Reduviid  bug,  Conorhiniis  rubrofasciattis. 
The  insect  attacks  man  in  Mauritius,  Reunion  and  other  places. 
Lafont  infected  rats  and  mice  by  intraperitoneal  injection  with  the 
gut-contents  of  infected  bugs.  Trypanosomes  appeared  in  the  mice. 
Other  flagellate  types  were  assumed  by  the  parasites  in  the  bug. 

MoNOMORPHic  Trypanosomes. 

A  number  of  trypanosomes,  characterized  by  relative  uniformity 
in  size  and  structure,  may  be  considered  under  this  heading.  They 
occur  in  cattle,  sheep,  goats  and  horses  in  Africa,  especially  West 
Africa.  Morphologically,  they  are  characterized  by  the  posterior 
(aflagellar)  part  of  the  body  being  swollen,  while  the  anterior  part 
narrows.  The  nucleus  is  central  and  situated  at  the  commencement 
of  the  narrowing  of  the  body.  The  blepharoplast  is  almost 
terminal,  the  undulating  membrane  is  narrow  and  not  markedly 
folded,  so  that  the  flagellar  border  lies  close  to  or  along  the  body. 
The  fiagellum  may  or  may  not  possess  a  free  portion. 

Some  recent  workers  have  considered  that  T.  briicel  (Zululand 
strain)  and  T.  evansl  are  also  monomorphic,  but  they  do  not  exhibit 
the  general  characteristics  outlined  above.  T.  hriicei  and  T.  evansi 
have  already  been  considered  separately. 

The  monomorphic  trypanosomes,  as  defined  above,  include  : — 

Trypanosoma  vivax,  Ziemann,  1905. 

This  trypanosome^  occurs  in  cattle,  sheep  and  goats,  and  was  first 
found  in  the  Cameroons.  It  is  fatal  to  cattle.  Equines  are  also 
affected.     Antelopes  are  the  possible  reservoirs  of  the  trypanosome. 

'  C.  R.  Soc.  Biol.y  Ixv,  p.  581.  2  Joitrn.  Ale.i.  Research^  xxxi,  p.  195. 

^  See  Bruce  and  colleagues  (1910),  Proc.  Roy.  Soc,  B,  Ixxxiii,  p.  15. 


TOO 


THE   ANIMAL   PARASITES   OF   MAN 


It  is  probably  transmitted  by  Glossina  palpalis  and  other  tsetse  flies. 
Its  movement  is  very  active.  It  possesses  a  free  flagellum  (fig.  45)  and 
it  averages  23//,  to  24  yit  in  length.  T.  cazalhoni  (Laveran,  1906) — the 
causal  agent  of  ''souma"  in  bovines  and  equines  in  the  French  Sudan 
— is  probablv  synonymous  with  T.  vlvax. 

Trypanosoma  caprae  (Kleine,  1910)  is  allied, 

but  is  somewhat   broader  and    more   massive. 

It  was  found  in  goats  in  Tanganyika. 

Trypanosoma  congolense,  Bi  oden,  1904. 

Probable  synonyms. —  Trypcmosoma  dimorphon,  Laveran 
and  Mesnil,  1904  ;  Trypa?iosofna  namun,  Laveran, 
1905  ;  Trypanosoma  pecorum^  Bruce,  1910  ;  Try- 
panosoma confusum,  Montgomery,  1909. 

This  trypanosome  causes  disease  among 
horses  {e.g.,  Gambia  horse  sickness),  cattle, 
sheep,  goats,  pigs,  and  dogs.  It  is  widely  dis- 
tributed in  Central  Africa  {e.g.,  Gambia,  Congo, 
Uganda,  Nyasaland),  the  strain  probably  being 
maintained  naturally  in  big  game.  It  is  trans- 
mitted by  various  Glossince,  and  perhaps  by  Tabanus  and  Stomoxys.  It 
is  said  to  develop  in  the  gut  and  proboscis  of  Glossina  palpalis  and 
G.  morsitans.  The  trypanosome  averages  13  /x  to  14  yLt  in  length  and 
has  no  free  flagellum  (fig.  46).  It  is  about  2  fj,  broad.  Formerly 
T.  tiaiium  and  T.  pecorumwere  said  to  differ  in  their  pathogenicity,  the 
former  being  said  not  to  infect  the  smaller  laboratory  animals.  Yorke 
and  Blacklock  (1913),  however,  consider 
that  the  virulence  varies  and  that  these 
trypanosomes  are  probably  the  same. 


Fig.  45.  —  Trypanosoma 
vivax.  x2,oco.  (Original. 
From  preparation  by  Fan- 
tham.) 


Fig.  46.  —  Trypanosoma 
congolense.  x  2,000.  (Orig- 
inal. From  preparation  by 
Fantham.) 


Fig.  47.  —  Trypanosoma 
uni forme,  x  2,000,  (Orig- 
inab  From  preparation  by 
Fantham.) 


The  T.  diinorphon  originally  obtained  by  Button  and  Todd  (1903) 
in  Gambian  horse  sickness  has  been  shown  to  be  a  mixture  of  T.  vivax 
and  T,  congolense. 

Trypanosoma  simiae  (7\  ignotum)  is  like  T.  congolense.  It  averages 
I7'5  /A  long.     It  is  virulent  to  monkeys  and  pigs. 


TRYPANOSOMA    UNI  FORME 


lOI 


Trypanosoma  unlforme,  Bruce,  1910. 

This  trypanosome  was  found  in  oxen  in  Uganda/  It  can  be  in- 
oculated to  oxen,  goats  and  sheep,  but  is  refractory  to  dogs,  rats  and 
guinea-pigs.  It  has  been  found  in  antelopes.  It  resembles  T.  vivax, 
but  is  smaller  (fig.  47),  averaging  16  fju  in  length.  A  free  flagellum 
is  present.    It  is  transmitted  by  Glossincv. 

Many  other  trypanosomes  occur  in  mam- 
mals, while  birds,  reptiles,  amphibia  (fig.  48) 
and  fish  also  harbour  them.  The  discussion  of 
these  forms  does  not  come  within  the  scope 
of  the  present  work.  They  are  dealt  with 
in  Laveran  and  Mesnil's  ''Trypanosomes  et 
Trypanosomiases,"  2nd  edit.,  19 12. 

General  Note  on  Development  of 

Trypanosomes    in    GLOSSINA.  Yig.^^.—  Trypanosoma 

rotatoriufn,  from  blood   of 

Before  concluding  the  account  of  trypano-     a  frog,    x  1,400.    (After 

..  ,  r    ■     1  11  1      ii     J        Laveran  and  Mesnll.) 

somes,   it  may   be  of  mterest  to   remark   that 

several  African  trypanosomes  develop  in  various  species  of  Glosshia, 
and  are  found  in  different  parts  of  the  alimentary  tract  and  in  the 
proboscis.  Thus  {a)  T.  vivax,  T.  uniforme  and  T.  caprce  develop  in  the 
fly's  proboscis  (labial  cavity  and  hypopharynx)  only ;  {b)  T.  congolense, 
T,  simicB  and  T.  pecaudi  develop  first  in  the  gut  of  the  fly  and  then 
pass  forward  to  its  proboscis ;  and  (c)  T.  gambiense  and  T.  rhodesiense 
develop  first  in  the  gut  and  later  invade  the  salivary  glands  of  the 
tsetse.  The  proboscis  or  the  salivary  glands  in  such  cases  are  termed 
by  Duke^  the  anterior  station  of  the  trypanosome,  wherein  it  completes 
its  development. 

Adaptation  of  Trypanosomes. 

These  flagellates  may  exhibit  power  of  adaptation  to  changes  of 
environment,  such  as  those  due  to  the  administration  of  drugs,  change 
of  host,  etc.  A  few  examples  of  such  mutations  may  be  briefly 
considered  : — 

(i)  Blepharoplastless  Trypanosomes. — T.  briicei  may  become  resistant 
to  pyronin  and  oxazine.  Accompanying  this  drug  resistance  is  a 
change  in  morphology,  namely,  the  loss  of  the  blepharoplast 
(Werbitzki).^  A  race  or  strain  of  blepharoplastless  trypanosomes  may 
be  thus  produced  which  retains  its  characteristic  feature  after  as  many 
as  130  passages  (Laveran).^  Oxazine  is  the  more  powerful  drug,  and 
it  acts  directly  on  the  blepharoplast.  (Compare  the  natural  blepha- 
roplastless character  of  T.  equintun.) 

'  Froc.  Roy.  Soc,  B;  Ixxxiii,  p.  176. 

2  Repts.  Sleeping  Sickness  Commission  Roy.  Soc.  (1913),  xiii,  p.  82. 

3  Centralbl.  f.  Bakt.  (1910),  Orig.,  liii,  p.  303.  *  Bull.  Soc.  Path.  Exot.,  iv,  p.  233. 


102  THE   ANIMAL   PARASITES   OF   MAN 

(2)  Reference  has  been  made  on  p.  93  to  the  experiments  of 
Gonder,  who  showed  that  a  strain  of  T.  lewisi  rendered  resistant  to 
arsenophenylglycin  lost  its  resistance  after  passage  through  the  rat 
louse.  This  is  in  marked  contrast  with  the  retention  of  drug  resistance 
during  passage  b}^  inoculation  from  rat  to  rat. 

(3)  T.  lewisi  from  the  blood  of  a  rat  when  transferred  to  a  snake 
seems  largely  to  disappear,  as  very  few  flagellates  are  seen.  When 
blood  from  the  snake  is  inoculated  into  a  clean  rat,  then  trypano- 
somes  reappear  in  the  rat,  but  they  are  not  all  like  those  originally 
inoculated.  It  seems  certain  that,  in  such  a  case,  changes  in  form 
and  virulence  of  the  trypanosome  have  occurred.  Similar  experiments 
were  made  with  T.  hriicei  from  rats  to  adders  and  other  animals  and 
back  to  rats.    Changes  in  the  form  and  virulence  of  T.  hrucei  occurred. 

These  interesting  experiments  were  performed  by  Wendelstadt  and 
Fellmer.' 

Genus.     Herpetomonas,  Saville  K'^nt,  1881. 

Herpetomonas  is  a  generic  name  for  certain  flagellates  possessing 
a  vermiform  or  snake-like  body,  a  nucleus  placed  approximately  cen- 
trally, and  a  blepharoplast  (kinetic  nucleus)  near  the  flagellar  end. 
There  is  no  undulating  membrane  (fig.  49,  a).  The  organisms  in- 
cluded in  this  genus  certainly  possess  one  flagellum,  while  according 
to  Prowazek  (1904)  Herpetomonas  miiscce-domesticcey  the  type  species, 
possesses  two  flagella  united  by  a  membrane.  Patton,^  Porter  ^  and 
others  affirm,  however,  that  the  biflagellate  character  of  H.  niiiscce- 
domesticcB  (from  the  gut  of  the  house-fly)  is  merely  due  to  precocious 
division.  The  matter  is  further  complicated  by  the  generic  name 
Leptomonas,  given  by  Kent  in  1881,  to  an  uniflagellate  organism  found 
by  Butschli  in  the  intestine  of  the  Nematode  worm,  Trilohus  gracilis. 
This  parasite,  Leptomonas  hiltschlii^  has  not  yet  been  completely 
studied.  Until  these  controversial  points  relating  to  the  identity  or 
separation  of  Herpetomonas  and  Leptomonas  have  been  satisfactorily 
settled,  we  may  retain  the  better  known  name  Herpetomonas  for  such 
uniflagellate,  vermiform  organisms.  However,  the  name  Leptomonas, 
having  been  used  by  Kent  two  pages  earlier  in  his  book  {^'  Manual  of 
the  Infusoria  ")  than  Herpetomonas,  would  have  priority  if  the  two 
generic  names  were  ultimately  shown  to  be  synonymous. 

A  full  discussion  of  these  interesting  and  important  flagellates 
hardly  comes  within  the  purview  of  the  present  work ;  brief  mention 
can  only  be  given  here  to  certain  species. 

The  Herpetomonads  occur  principally  in  the  digestive  tracts  of 
insects,  such  as  Diptera  and  Hemiptera.     They  are  also  known  in  the 

'  Zeitschr.f.  Immunitatsforschung^  iv,  p.  422  (1909),  and  v,  p.  337  (1910). 
*  Arch.f.  Protist.^  xiii,  p.  i.  ^  Parasitology^  ii,  p.  367. 


HERPETOMONAS 


103 


guts  of  fleas  and  lice,  but  are  not  confined  to  blood-sucking  insects. 
One  example,  H.  deuocephali  (Fantham,  191 2) '  occurs  in  the  digestive 
tracts  of  dog  fleas,  Ctenocephalns  canis,  in  England,  France,  Germany, 
Italy,  India,  Tunis,  etc.  It  is  a  natural  flagellate  of  the  flea,  and  might 
easily  be  confused  with  stages  of  blood  parasites  in  the  gut  of  the  dog 
flea.  Dog  fleas  are  stated  by  Basile  to  transmit  canine  kala-azar,  which 
is  believed  to  be  the  same  as  human  infantile  kala-azar.  Confusion  is 
further  likely  to  arise  since  herpetomonads  pass  through  pre-flagellate, 
flagellate  and  post-flagellate  or  encysted  stages;  pre-  and  post-flagellate 
stages  being  oval  or  rounded  and  Leishmania-like.  The  post-flagellate 
stages  are  shed  in  the  faeces,  and  are  the  cross-infective  stages  by 
means  of  which  new  hosts  are  infected  by  the  mouth.  The  possible 
presence  of  such  natural  flagellates  must  alw^ays  be  considered  when 
experimenting  with  fleas, 
lice,  mosquitoes,  etc.,  as 
possible  vectors  of  patho- 
genic flagellates  like  Leish- 
mania  and  Trypanosoma, 
H,  pedicidi  (Fantham, 
191 2)  occurs  in  human 
body  lice.^  See  further 
remarks  on  pp.  107,  112. 

Laveran  and  Franchini 
(1913-14)^  have  recently 
succeeded  in  inoculating 
Herpetoinonas  ctenocepJiali, 
from  the  gut  of  the  dog 
flea,  intraperitoneally  into 
white  mice,  and  produc- 
ing an  experimental  leish- 
maniasis in  the  mice.     A 

dog  was  also  infected.  They  have  also  succeeded  in  infecting  mice 
with  H.  pattoni — a  natural  flagellate  of  the  rat  flea — by  mixing  in- 
fected rat  fleas  with  the  food  of  the  mice,  and  by  causing  them  to 
ingest  infected  faeces  of  rat  fleas.  Further,  they  have  shown  that 
infection  with  the  herpetomonas  occurs  naturally  by  this  method, 
that  is,  by  the  rodents  eating  the  fleas  and  not  by  the  insects  inocu- 
lating the  flagellates  into  the  vertebrates  when  sucking  blood.  These 
experiments  shed  an  interesting  light  on  the  probable  origin  of 
Leishmania  and  its  cultural  herpetomonad  stage,  which  were  very 
probably  once  parasitic  flagellates  in  the  gut  of   an  insect. 

•  Bull.  Path.  Exot.,  vi,  p.  254.  ^  Froc.  Roy.  Soc,  B,  Ixxxiv,  p.  505. 

^  C.  R.  Acad.  Set.,  clvii,   pp.   423,   744.     Ibid.,  clviii,    pp.   450,  770.     Bull.  Soc.  Paih. 
Exot.,  vii,  605. 


Fig.  49. — a,  Herpetomonas  ;  b,  Crithidia 
Cy  Trypanosoma.     (After  Porter.) 


104  THE   ANIMAL   PARASITES   OF   MAN 

Fantham  and  Porter^  (1914-15)  have  shown  that  young  mice  may 
be  inoculated  or  fed  with  Herpetomonas  jaculum,  from  the  gut  of  the 
Hemipteran,  Nepa  ciiierea  (the  so-called  "water-scorpion"),  with  fatal 
results.  The  pathogenic  effects  are  like  those  of  kala-azar.  They  also 
showed  that  the  post-flagellate  stages  of  tlie  herpetomonads  seemed 
most  capable  of  developing  in  the  vertebrate. 

A  herpetomonad,  N.  davidi^  has  been  found  in  the  latex  of  species  of  the  plant- 
genus  Euphorbia  in  Mauritius,  India,  Portugal,  etc.  It  is  apparently  transmitted  to 
the  plants  by  Hemiptera.     The  plants  sometimes  suffer  from  "  flagellosis." 

Franchini  (1913)  ^  has  described  a  new  parasite,  Hccnwcystozoon 
brasiliense,  from  the  blood  of  a  man  who  had  lived  in  Brazil  for  many 
years.  It  possesses  flagellate  and  rounded  stages,  and  is  closely  allied 
to  the  herpetomonads. 

Genus.     Crithidia,  Leger,  1302,  emend.  Patton,  1908. 

Crithidia  is  the  generic  name  of  vermiform  flagellates  with  a 
central  nucleus,  a  blepharoplast  or  kinetic  nucleus  in  the  neighbour- 
hood of  the  principal  nucleus,  and  a  rudimentary  undulating  mem- 
brane bordered  by  a  flagellum  arising  from  a  basal  granule,  which  is  the 
centrosome  of  the  kinetic  nucleus  (fig.  49^).  The  anterior  or  flagellar 
end  of  the  body  is  attenuated  and  fades  off  as  the  undulating  membrane. 

Crithidia  fasciculata,  the  type  species,  was  found  by  Leger  in  the 
alimentary  canal  of  Anopheles  macidipennis.  Crithidia  occur  in  bugs, 
flies,  fleas,'  and  ticks.  Some  of  them  are  found  in  the  body-fluid 
of  the  invertebrate  host  as  well  as  in  the  gut.  Others  may  be  restricted 
to  the  body  cavity  or  intestine  respectively.  C.  melopliagia  from  the 
sheep-ked,  Melophagns  ovintis,  and  C.  hyaloinmce  from  the  haemocoelic 
fluid  of  the  tick,  Hyalotmna  cegyptiutUj  pass  into  the  ovaries  and  eggs 
of  their  hosts,  and  the  young  keds  or  ticks  are  born  infected. 

C.  fasciciilata  has  been  shown  by  Laveran  and  Franchini  to  be 
inoculable  into  white  mice,  producing  a  sort  of  experimental  leish- 
maniasis therein.  In  one  case  cutaneous  lesions  were  produced  like 
those  of  Oriental  sore. 

Crithidia  are  natural  flagellates  of  Arthropoda,  with  their  own 
pre-flagellate,  flagellate  and  post- flagellate  stages,  and  must  not  be 
confused  with  transitory  crithidial  stages  of  trypanosomes. 

Genus.     Leishmania,  Ross,  1903. 

With  an  oval  body  containing  nucleus  and  blepharoplast  (kinetic  nu- 
cleus) but  no  flagellum.    An  intracellular  parasite  in  the  vertebrate  host. 
Included  in  the  genus  Leishmania  are  three  species,  namely  : — 

'  Froc.  Camb.  Philosoph.  Soc,  xviii,  p.  39.     "■  BtilL  Soc.  Path.  Exot.,  vi,  pp.  156,  333,  377. 
^  See  Porter,  Parasitology,  iv,  p.  237. 


LEISHMANIA    DONOVANI  105 

(i)  Leishmania  donovaiii,  Laveran  and  Mesnil,  1903,  the  parasite 
of  Indian  kala-azar,  a  generalized  systemic  disease,  usually 
fatal,  occurring  in  subjects  of  all  ages. 

(2)  Leislunania  tropica,  Wright,   1903,  the  parasite  of  Delhi  boil, 

Oriental  sore,  Aleppo  button — a  localized,  cutaneous  disease, 
usually  benign. 

(3)  Leishmania  infantujii,  Nicolle,   1908,  the  parasite   of   infantile 

kala-azar,  occurring  in  children  (and  a  few  adults)  around 
the  shores  of  the  Mediterranean.  The  disease  is  perhaps  a 
form  of  Indian  kala-azar,  and  the  parasite  is  probably  identical 
with  L.  donovani. 
These  diseases  may  be  termed  collectively  leishmaniases.  The 
morphology  of  the  various  species  is  practically  identical. 

Leishmania  donovani,  Laveran  and  Mesnil,  1903. 
Syn.  :  Piroplasjna  donovam\  Laveran  and  Mesnil. 

The  parasite  of  Indian  kala-azar  was  demonstrated  in  1900  by 
Leishman  from  a  post-mortem  examination  of  a  case  of  ^*  Dum-Dum 
fever,"  but  details  were  not  published  till  May,  1903.  In  July,  1903, 
Donovan  found  similar  bodies  from  cases  in  Madras.  Rogers 
succeeded  in  cultivating  the  parasite  in  July,  1904.^  The  original 
centre  of  the  disease  was  probably  Assam  ;  it  occurs  also  in  Madras, 
Ceylon,  Burma,  Indo-China,  China  and  Syria.  A  variety  of  this  leish- 
maniasis is  found  in  the  Sudan.  The  patient  becomes  emaciated, 
with  a  greatly  enlarged  spleen.     There  is  anaemia  and  leucopenia. 

The  parasite,  commonly  known  as  the  Leishman-Donovan  body^ 
is  intracellular  (fig.  50,  2,  5).  It  is  found  in  the  endothelial  cells  of  the 
capillaries  of  the  liver,  spleen,  bone-marrow,  lymphatic  glands  and 
intestinal  mucosa,  and  in  the  macrophages  of  the  spleen  and  bone- 
marrow.  Some  host  cells  may  contain  many  parasites.  It  is  rather 
rare  in  the  circulating  blood,  but  may  be  found  in  the  blood  from 
the  femoral,  portal  and  hepatic  veins.  It  does  not  occur  in  the  red 
blood  corpuscles  as  was  formerly  thought.  The  parasites  liberated 
from  the  endothelial  cells  are  taken  up  by  the  mononuclear  and 
polymorphonuclear  leucocytes.  The  Leishman-Donovan  body  is  the 
resting  stage  of  a  flagellate.  As  found  in  man  it  is  a  small,  oval 
organism,  about  2*5 //,  to  3*5//'  in  length  by  2  yu,  in  breadth,  and  con- 
taining two  chromatinic  bodies,  corresponding  to  the  nucleus  and 
kinetic  nucleus  (blepharoplast)  of  a  flagellate.  The  latter  element  is 
the  smaller  and  more  deeply  staining,  and  is  usually  placed  at  the 
periphery,    transversely   to    the    longer   axis   of   the    oval    organism. 

^  The  literature  up  to  191 2,   on    kala-azar   and   other   leishmaniases   is   reviewed   in    the 
Kala-azar  Bulletin.     Afterwards  in  the  Tropical  Diseases  Bulletin. 


io6 


THE   ANIMAL   PARASITES   OF   MAN 


There  is  sometimes  a  very  short,  slightly  curved  filament  to  be  seen,, 
which  may  be  a  rhizoplast.  Multiplication  takes  place  by  binary  or 
multiple  fission.  The  presence  of  the  parasite  used  to  be  demonstrated 
by  splenic  or  hepatic  prmcture  ;  nowadays  it  can  be  demonstrated  in. 
peripheral  blood,  e.g.,  of  the  finger,  or  by  culture  of  infected  blood. 


Fig.  50. — Leishmania  donovani.  /,  't  lee  lorms,  each  with  nucleus  and  rod -shaped  Mepharo- 
plast  (after  Christophers)  ;  2,  endothelial  cell  and  leucocytes  containing  parasites  (after 
Christophers) ;  j,  capillary  in  the  liver  showing  endothelial  cells  containing  parasites  (after 
Christophers)  ;  ^,  two  parasites  escaping  from  a  leucocyte  in  the  alimentary  canal  of  the  bug 
(after  Palton)  ;  5,  further  development  in  bug  (after  Patton)  ;  6,  young  flagellate  forms  in  bug 
(after  Patton) ;  7-/7,  culture  forms  (after  Leishman)  ;  7,  8^  9,  show  development  of  flagellum. 

L.  donovani  can  be  cultivated  in  citrated  splenic  blood,  under 
aerobic  conditions,  at  22°  to  25°  C.  This  was  first  accomplished 
by  Rogers  (1904).  It  is  not  so  easily  culturable  as  L.  infantum  on 
the  Novy-MacNeal-Nicolle  medium.^     L.  donovani  is  inoculable  with 

'  For  the  composition  of  this  medium,  see  Appendix. 


LEISHMANIA   TROPICA  107 

some  difficulty  into  experimental  animals — in  India,  white  rats,  white 
mice,  dogs  and  monkeys  (Macaais  spp.),  have  been  inoculated.  The 
Sudan  variety,  somewhat  less  virulent,  is  inoculable  to  monkeys.  Row 
also  produced  a  local  lesion  in  Macaais  slnicus  by  subcutaneous  inocu- 
lation of  L.  dofiovani.  Parasites  taken  from  such  a  local  lesion  were 
found  to  be  capable  of  producing  a  generalised  infection  in  Macacus 
slnicus  and  white  mice. 

In  cultures  the  various  species  of  Leishmania  all  grow  into 
herpetomonad,  uniflagellate  organisms  (fig.  50,  10),  about  12  fi  io  20  /j, 
in  body  length.  On  this  account  Rogers^  and  Patton  place  the 
Leishman-Donovan  body  within  the  genus  Herpetoinonas.  The 
method  of  culture  may  be  used  in  diagnosing  leishmaniases. 

Kala-azar  is  very  probably  an  insect- borne  disease.  Patton^  sus- 
pects the  bed-bug  to  be  the  transmitter  and  finds  (fig.  50,  4-6)  that  the 
Leishman-Donovan  body  can  develop  mto  the  flagellate  stage  in  the 
digestive  tract  of  the  bed-bug.  Feeding  experiments  are  unsatisfactory, 
since  there  are  very  few  cases  in  which  the  parasites  occur  in  sufficient 
numbers  in  the  peripheral  blood  to  make  the  infection  of  the  insect 
possible,  or  at  any  rate  easy.  In  examining  the  alimentary  tracts  of 
insects  for  possible  flagellate  stages  of  Leishmania,  it  must  be  remem- 
bered that  in  many  insects  natural  flagellate  parasites,  belonging  to  the 
genus  Herpetontonas,  m3.y  occur  therein  ;  such  natural  insect  flagellates 
may  be  harmless,  and  have  no  connection  with  the  life-cycle  of 
L.  donovani.  Natural  herpetomonads  are  known  to  occur  in  the 
alimentary  tracts  of  flies,  mosquitoes,  sand-flies,  fleas  and  lice,  but 
not  in  bed-bugs.  Further,  if  such  flagellates  are  able  to  be  inoculated 
into  and  live  within  vertebrate  hosts,  producing  symptoms  like  those 
of  leishmaniasis,  the  origin  of  kala-azar  is  indicated  (see  pp.  104,  112). 

Leishmania  tropica,  Wright,  1903. 

Syn.  :  Helcosoma  tropicum,  Wright,  1903  ;  Z.  wrighti,  Nicolle,  1908;  Ovoplasma 
orientale^  Marzinowsky  and  Bogrow. 

It  is  believed  by  some  that  the  parasite  was  first  described  by  Cunningham  in 
1885,  and  studied  by  Firth  in  1891,  being  called  by  him  Sporozoon  furuncidosum. 
If  these  earlier  studies  were  of  the  parasite,  then  its  correct  name  is  L.  furunculosa^ 
Firth,  1891. 

The  benign  disease  produced  by  this  parasite  has  received  many 
names,  among  the  best  known  being  Oriental  sore.  Tropical  sore, 
Delhi  boil  and  Aleppo  button.  These  names,  however,  are  not  happy 
ones,  as  cutaneous  leishmaniasis  {e.g.,  on  the  ear)  is  now  known  to 
occur  in  the  New  World,  for  example  in  Mexico,  Venezuela,  Brazil 
and  neighbouring  States.  However,  it  may  be  necessary  to  subdivide 
cutaneous  leishmaniases  later. 

'  Proc.  Roy.  Soc,  B,  Ixxvii,  p.  284.  2  Sci.  Mem,  Govt.  India,  Nos.  27,  31  (1907-08). 


I08  THE   ANIMAL   PARASITES   OF   MAN 

In  the  Old  World  the  disease  occurs  in  India,  Persia,  Arabia  and 
Transcaucasia.  It  is  also  known  in  Algeria,  Northern  Nigeria,  Egypt, 
Sudan,  Crete,  Calabria,  Sicily  and  Greece. 

The  boils  often  occur  on  the  face,  and  before  ulceration  the 
parasites  may  be  found  in  the  cells  at  the  margin  and  floor  of  the 
"  button."  In  searching  for  parasites  the  scab  should  be  removed  and 
scrapings  made  from  the  floor  and  edges.  Where  lesions  occur 
atrophy  of  the  epidermis  takes  place,  and  infiltration  of  mononuclear 
cells  (e.g.,  plasma  cells,  lymphoid  and  endothelial  cells)  follows. 
The  parasites  are  intracellular,  being  found  inside  mononuclear  cells. 
In  non-ulcerating  sores,  Cardamitis  found  some  free  parasites.  Non- 
ulcerating  forms  are  said  to  occur  in  the  Sudan.  In  the  Old  World 
the  sores  are  often  limited  to  exposed  surfaces  of  the  body.  Infection 
of  mucous  membranes  (such  as  the  lip,  palate,  buccal  and  nasal 
membranes)  may  occur,  especially  in  South  America,  and  are  often 
known  there  as  ^^  Espundia."  Christopherson  (1914)  has  recorded  a 
case  in  Khartoum. 

Leishmania  tropica  is  equally  well  cultivated  on  Novy-MacNeal- 
Nicolle  medium  or  on  citrated  blood.  The  usual  temperature  for 
cultivation  is  22°  to  28°  C,  though  Marzinowski  claims  to  have 
cultivated  the  parasite  at  37°  C.  L.  tropica  can  be  inoculated 
into  monkeys  and  dogs,  with  the  production  of  local  lesions. 
Material  from  a  human  sore  or  flagellates  from  a  culture  may  be  thus 
successfully  inoculated.  Also  infected  material  may  be  rubbed 
directly  into  a  scarified  surface.  The  incubation  period  is  long, 
extending  over  several  months.  The  duration  of  the  disease  may  be 
from  twelve  to  eighteen  months.  Recovery  from  one  attack  of  tropical 
sore  confers  immunity,  and  the  Jews  in  Bagdad  inoculate  their  children 
with  the  disease  on  a  part  of  the  body  which  will  be  covered,  and 
so  secure  immunity  in  adult  life. 

The  mode  of  transmission  of  L.  tropica  is  unknown.  Wenyon 
(1911)^  has  found  that  the  parasite  develops  into  the  flagellate  stage 
in  the  digestive  tract  of  Stegomyia  fasciata  in  Bagdad.  Patton  (191 2)' 
has  found  similar  development  in  the  bed-bug  in  Cambay.  The 
house-fly,  Phlehotonins  and  Simulium  have  been  suspected  as  trans- 
mitters in  different  parts  of  the  world. 

An  interesting  announcement  has  been  made  recently  (May,  1913), 
that  Neligan  has  found  that  L.  tropica  occurs  in  dogs  in  Teheran, 
Persia,  producing  ulcers  on  the  dogs'  faces  [cf.  natural  occurrence  of 
L.  infantiim  in  dogs — see  p.  no).  Yakimoff  and  Schokhor  (1914),^ 
have  found  the  disease  in  dogs  in  Tashkent. 

Gonder*    (191 3)     has     performed    some    interesting    experiments 

'  Parasitology,  iv,  p.  387.  '  Sci.  Mem.  Govt.  India,  No.  50. 

3  Bull.  Sue.  Path.  Exot.,  vii,  p.  i86-.  *  Arch.  f.  Schifs-  u.   Prop..  Hyg.^  xvii,  p.  397. 


LEISHMANIA   INFANTUM 


109 


showing  the  relation  of  infantile  kala-azar  to  Oriental  sore.  Gonder 
infected  mice  with  L.  infantum  and  with  L.  tropica.  He  used  culture 
material  and  injected  intraperitoneally  or  intravenously.  In  each  a 
general  infection  resulted,  with  enlargement  of  the  liver  and  spleen. 
Later,  however,  mice  injected  with  Oriental  sore  (North  African 
variety)  developed  peripheral  lesions  on  the  feet,  tail  and  head,  and 
the  lesions  contained  Leishuiania.  No  such  peripheral  lesions 
developed  in  the  case  of  the  mice  infected  with  the  kala-azar  virus. 
Gonder  suggested  that  Oriental  sore,  like  kala-azar,  is  really  a  general 
infection  overlooked  in  its  earlier  stages,  and  that  it  is  in  the  later 
stages  that  peripheral  lesions  on  the  skin  are  developed.  Row  (1914)^ 
also  obtained  a  general  infection  in  a  mouse  by  the  injection  of 
cultures  of  L.  tropica  from  Oriental  sore  of  Cambay. 

Leishmania  Infantum,  Nicolle,  1908.^ 

Infantile  splenic  anaemia  has  been  long  known  in  Italy.  It  also 
occurs  in  Algeria,  Tunis,  Tripoli,  Syria,  Greece,  Turkey,  Crete,  Sicily, 
Malta,^  Spain  and  Portugal.  This  leishmaniasis  is, then, distributed  along 
the  Mediterranean  littoral;  also  in  Russia.  Cathoire  (1904)  in  Tunis 
and  Pianese  (1905)  in  Italy  were  among  the  first  to  see  the  parasite. 
Nicolle  then  found  the  parasite  in  patients  in  Tunis,  and  further  found 
spontaneous  infection  in  dogs.  The  patients  are  usually  children 
between  the  ages  of  2  and  5  years.  There  are  a  few  cases  known  in 
which  the  infantile  type  of  leishmaniasis  occurred  in  youths  and  adults 
of  the  ages  of  17  to  19,  while  one  patient  in  Calabria  was  38  years  old. 
The  symptoms  are  like  those  of  Indian  kala-azar.  Several  Italian 
investigators  and  others  consider  that  L.  infantum  is  the  same  as 
L.  donovani,  and  that  the  latter  name  should  be  used  for  the  parasite 
of  Mediterranean  leishmaniasis.  This  view,  as  to  the  identity  of 
L.  donovani  and  L.  infantum,  seems  coming  into  general  favour. 

There  are,  however,  differences  between  the  Indian  and  infantile 
kala-azars,  in  addition  to  the  ages  of  the  patients  affected,  thus  : 
{a)  As  regards  cultures,  it  is  found  that  L.  infantum  is  readily  grown 
on  the  Novy-MacNeal-Nicolle  C'  N.N.N.")  medium  (saline  blood- 
agar),  and  that  sub-cultures  are  easily  obtained ;  in  citrated  blood 
L.  infantum  grows  with  difficulty.  The  reverse  is  the  case  with 
regard  to  culture  media  for  L.  donovani,  which  grows  with  difficulty 
on  the  N.N.N,  medium,  but  relatively  easily  in  citrated  splenic  blood. 
(6)  Considering  inoculability  into  experimental  animals,  it  is  found  that 
L.  donovani  is  inoculated  generally  with  some  difficulty  into  white  rats, 

*  Bull.  Soc.  Path.  Exot.^  vii,  p.  272.  ^  Arch.  Inst.  Pasteur  Tunis,  i,  p.  26. 

^  See  Wenyon  (1914),    Trans.  Soc.   Trop.  Med.  attd  Hyg.,  vii,  p.  97;  also  Critien  (191 1), 
Annals  Trop.  Med.  and  Parasitol.,  v,  p.  37. 


no  THE   ANIMAL   PARASITES   OF   MAN 

white  mice  and  monkeys,  and  with  greater  difficulty  into  dogs,  while 
L.  infaninm  can  be  inoculated  into  several  experimental  animals, 
especially  into  dogs  and  monkeys,  with  ease,  (c)  At  present  L.  donovani 
is  not  known  to  occur  spontaneously  in  animals,  jDut  L.  infantiun  is 
found  naturally  in  dogs  in  the  Mediterranean  region,  and  the  disease  in 
dogs  is  often  referred  to  as  canine  kala-azar.  Kittens  have  occasionally 
been  found  infected.  However,  these  differences  must  not  be  empha- 
sized too  much. 

The  material  for  cultivation  is  obtained  from  punctures  of  spleen, 
liver  or  bone-marrow  of  cases  infected  with  L.  infantum.  It  is  not 
always  easy,  however,  to  infect  from  cultures,  as  the  cultural  flagellates 
inoculated  into  the  body  are  often  phagocytosed. 

Similarly,  the  material  for  animal  inoculation  is  obtained  from 
emulsions  of  infected  spleen,  liver  or  bone-marrow.  Dogs  and 
monkeys  are  easily  inoculated  with  such  material ;  Nicolle  inoculates 
into  the  liver  or  the  peritoneal  cavity.  Mice,  white  rats,  guinea-pigs 
and  rabbits  only  show  slight  infections  after  such  inoculations. 

Dogs  infected  experimentally  with  infantile  leishmaniasis  may 
show  either  acute  or  chronic  symptoms.  The  acute  course  occurs 
more  often  in  young  dogs,  and  is  usually  fatal  in  three  to  five  months. 
The  chronic  course  is  found  more  commonly  in  older  dogs,  and  may 
last  seventeen  to  eighteen  months.  In  acute  forms  there  is  irregular 
fever,  progressive  wasting,  diarrhoea  occasionally,  motor  disturbances 
involving  the  hind  quarters,  and  the  animal  dies  in  a  comatose  con- 
dition. In  the  chronic  form  the  animal  may  appear  well,  except  for 
loss  of  weight.  The  parasites  may  be  found  in  the  internal  organs  of 
these  experimental  dogs,  but  are  not  numerous  in  the  peripheral 
blood  except  at  times  of  high  fever.  Experimental  monkeys  live  about 
three  months. 

It  may  be  interesting  to  record  the  number  of  dogs  found  to  be 
infected  naturally  with  leishmaniasis  in  various  countries.  In  Tunis, 
Nicolle  and  Yakimoff  found  about  2  per  cent,  infected  out  of  about 
500  dogs  examined.  Sergent  in  Algiers  found  9  infected  out  of  125 
dogs  examined.  In  Italy  and  Sicily,  Basile  found  about  40  per  cent, 
of  the  dogs  to  be  infected  out  of  93  examined  at  Rome  and 
Bordonaro.  Cardamitis  found  15  infected  out  of  184  examined  in 
Athens.  In  Malta,  Critien  found  3  infected  out  of  30  dogs  examined. 
Alvares  found  i  infected  dog  out  of  19  examined  in  Lisbon.  Pringault 
has  recently  (December,  191 3)  found  an  infected  dog  in  Marseilles.^ 
Yakimoff  and  Schokhor  found  24  per  cent,  infected  out  of  647  dogs 
examined  in  Turkestan. 

The  distribution  of  the  parasites  in  the  body  of  the  human  patient 
is  much  the  same  as  in  the  case  of  Indian  kala-azar.     Critien  records 


Bull.  Soc.  Path.  Exot.. 


p.  41 


LEISHMANIA   INFANTUM  IH 

the  finding  of  parasites  in  the  mucous  flakes  of  the  stools  of  a  three- 
year-old  Maltese  child/  Intestinal  lesions  rarely  occur  in  infantile 
leishmaniasis. 

Etiology. — Infantile  leishmaniasis  is  stated  to  be  transmitted  by 
fleas,  especially  dog  fleas,  Ctenocephalus  cams  (=  Pulex  serraticeps),  and 
by  Pulex  irritans.  Children  living  in  contact  with  infected  dogs  may  be 
bitten  by  infected  dog  fleas,  and  so  contract  the  disease.  Basile  (1910-11) 
and  Sangiorgi  (1910)  state  that  they  found  L.  infantum  parasites  in  the 
digestive  tract  of  the  dog  flea.  After  searching  they  found  infected 
dog  fleas  on  the  beds,  mattresses,  and  pillows  used  by  children 
suffering  from  the  disease.  Franchini  (191 2)  thinks  that  Anopheles 
maculipennis  may  be  concerned  in  the  transmission. 

Basile^  tried  a  number  of  experiments  to  show  that  infantile 
leishmaniasis  is  transmitted  by  fleas,  thus  : — 

(i)  Fleas  were  taken  from  a  healthy  dog.  They  were  placed  in 
vessels  containing  infected  spleen-pulp  and  allowed  to  feed  thereon. 
The  fleas  were  then  killed  and  dissected,  and  portions  of  the  gut- 
contents  examined  for  parasites.  The  remainder  of  the  gut  was 
emulsified  and  injected  into  a  young  puppy,  whose  bone-marrow  had 
been  shown  previously  to  be  uninfected.  Basile  states  that  the  puppy 
became  infected.  The  parasites  are  said  to  increase  in  number  in 
the  flea's  gut. 

(2)  Two  healthy  pups,  each  a  month  old,  and  born  in  the  labora- 
tory, were  placed  in  a  disinfected,  flea-proof  cage.  A  few  days  after, 
an  infected  dog  was  placed  in  the  cage,  so  that  fleas  from  the 
infected  dog  could  pass  on  to  the  puppies.  A  month  later  the  two 
pups  became  infected,  parasites  being  found  in  them  after  liver 
puncture.  A  number  of  control  puppies  from  the  same  litter 
remained  uninfected  and  in  good  health. 

(3)  Basile  next  used  other  laboratory-born  puppies,  a  month  old. 
Four  of  the  litter  were  placed  in  a  disinfected,  flea-proof  gauze  cage 
in  Rome.  The  cage  was  isolated  from  other  dogs.  Fleas  obtained 
from  an  infected  area  in  Sicily  were  placed  in  the  cage.  The  puppies 
were  examined  by  hepatic  puncture,  but  w^ere  found  to  be  negative 
for  two  months.  Then  two  of  the  puppies  showed  infection,  and 
six  days  later  the  remaining  two  puppies  were  found  to  be  infected, 
and  all  four  died.  They  showed  irregular  temperatures,  and  were 
getting  thin.     Control  puppies  remained  healthy. 

From  these  experiments  Basile  concludes  that  fleas  transmit 
leishmaniasis.  However,  Basile  did  not  exclude  the  possible  occur- 
rence of  natural  herpetomonads  in  the  gut  of  the  fleas.^    Herpetomonas 

'  Quoted  by  Leishman  (191 1)  in  his  interesting  review  of  Leishmaniasis,  lourn.  Foy.  Army 
Med.  Corps,  xvii,  p.  567,  xviii,  pp.  I,  125.     Also  Quart,  fourn.  Med.  v,  pp.  109-152. 
"^  Numerous  papers  in  Rendiconti  R.  Accad.  dei  Lincei  (Rome),  xix,  xx  (1910-II). 
8  See  Fantham,  Brit.  Med.  foiirn.^  1912,  ii,  p.  1 196. 


112  THE   ANIMAL   PARASITES   OF  MAN 

ctenocephali  is  known  to  occur  in  the  gut  of  Ctenocephalus  canis 
A  natural  Herpetoinonas  is  also  known  in  the  gut  of  P^ilex  irritans, 
as  well  as  a  Crithidla  (C.  piilicis,  Porter).  These  natural  flagellates  of 
the  fleas  pass  through  non-flagellate  stages,  like  the  Leishman-Donovan 
body.  In  consequence  Wenyon  and  Patton,  among  others,  have 
criticized  Basile's  results.  Further,  other  investigators,  such  as 
Wenyon  and  Da  Silva  (191 3),  have  repeated  Basile's  flea  experiments 
and  been  unable  to  confirm  them. 

In  feeding  and  inoculation  experiments  the  incubation  period  of 
the  parasite  may  be  long,  and  so  it  is  necessary  to  wait  a  long  time  to 
see  whether  the  parasite  will  develop. 

Iminnnity. — Nicolle  has  tried  some  experiments  with  L.  infantum 
and  L.  tropica.  He  finds  that  in  animals  recovery  from  an  attack  of  the 
former  confers  immunity  against  infection  by  the  latter  and  vice-versa. 

Laveran^  records  that  a  monkey  having  an  immunity  against  L. 
infantum  was  also  immune  to  L.  donovani. 

As  mentioned  on  p.  103,  Laveran  and  Franchini  (19 13),  working  in 
Paris,  have  succeeded  in  inoculating  Herpetomonas  ctenocephali,  a 
natural  flagellate  in  the  gut  of  the  flea,  Ctenocephalus  canis,  into 
white  mice.  Leishmaniform  stages  of  the  flea  flagellate  were  re- 
covered from  the  peritoneal  exudate,  blood  and  organs  of  the  mice 
some  weeks  after  inoculation.  The  parasites  may  also  be  conveyed  by 
way  of  the  digestive  tract  of  the  vertebrate.  Similar  experiments  have 
succeeded  with  H.  pattoni.  These  experiments  go  to  show,  together 
with  those  of  Fantham  and  Porter  with  H.  jaculum  (seep.  104),  that,  in 
the  words  of  the  latter  authors,  *^  it  may  be  expected  that  the  various 
leishmaniases,  occurring  in  different  parts  of  the  world,  will  prove  to 
be  insect-borne  herpetomoniases." 

Genus.     Histoplasma,  Darling,  1906. 

Under  the  name  Histoplasma  capsulattim,^  Darling  described  small 
round  or  oval  parasites,  enclosed  in  a  refractile  capsule,  and  each  con- 
taining a  single  nucleus.  The  bodies  were  found  in  cases  of  spleno- 
megaly in  Panama.  They  occurred  in  the  endothelial  cells  of  the 
small  blood-vessels  of  the  liver,  spleen,  lungs,  intestine  and  lymphatic 
glands,  and  also  within  the  leucocytes.  A  few  flagellates  were  stated  to 
occur  in  the  lungs.  The  parasite  has  usually  been  placed  near  Leish- 
mania,  but  recently  Rocha-Lima  has  stated  that  Histoplasma  is  a  yeast. 

Genus.     Toxoplasma,  Nicolle  and  Manceaux,  1908. 

The  genus  was  created  for  crescentic,  oval  or  reniform  parasites,  2*5  ju  to  6  /u  by 
2  Ai  to  3  /i,  possessing  a  single  nucleus  and  multiplying  by  binary  fission.     They  occur 

^  Annales  Inst.  Pasteur  (1914-15),  xxviii,  pp.  823,  885  ;  xxix,  pp.  i,  71. 

"^  lotirn.  Amer.  Med.  Assoc,  xlvi,  p.  1283  :  /ourn.  Exptl.  Med.  (1909),  xi,  p.  515. 


TOXOPLASMA 


113 


in  mononuclear  and  polymorphonuclear  cells  in  the  blood,  spleen,  liver,  peritoneum 
etc.  (fig.  51).  The  parasites  have  been  found  in  the  gondi,  dog,  rabbit,  mole,  mouse, 
pigeon  and  other  birds.  Although  various  species  names  have  been  given  to  the 
parasites  in  these  hosts,  it  seems  probable,  from  cross  infection  experiments,  that 
there  is  but  one  species  with  several  physiological  races.  Splendore'  (1913)  has 
described  a  flagellate  stage. 

Castellani  (1913-14)^  has  described  similar  parasites  from  a  case  of 
splenomegaly,  with  fever  of  long  standing,  in  a  Sinhalese  boy.     The 


Fig.  51. — Toxoplasma  gondii,  endocellular  or  free  in  the  peritoneal  exudate  of  infected 
mice.  I,  2,  mononuclear  leucocytes  containing  toxoplasms.  3,  polynuclear,  containing 
parasites.  4,  5,  6,  endothelial  cells  containing  toxoplasms,  agglomerated  in  6.  7,  agglomera- 
tion forms.    8-1 1,  free  forms.    12-13,  division  stages.     X  i,6do.    (After  Laveran  and  MaruUaz.) 


4» 


^^ 


■V 


9.    ^^ 


':i 


Fig.  ^2.— Toxoplasma  pyro^eius.  I,  body  found  in  blood.  2-7,  bodies  found  in  spleen, 
[i  is  about  the  size  of  a  red  blood  corpuscle,  as  drawn  in  the  figures!.  Magnification  not 
stated.     (After  Castellani.) 

bodies  were  found  in  the  spleen  and  more  rarely  in  the  blood  (fig.  52). 
Castellani  has  named  them  Toxoplasma  pyrogeiies.  Further  researches 
are  needed. 


^  Bull.  Soc.  Path.  ExoL,  vi,  p.  31 J 


■^  Journ.  Trop.  Med.  and  Hyg.,  xvii,  p.  113. 


114 


THE   ANIMAL   PARASITES   OF  MAN 


THE  SPIROCH.'^TES. 

The  Spirochaetes  are  long,  narrow,  wavy,  thread-like  organisms, 
with  a  firm  yet  flexible  outer  covering  or  periplast.  There  is  a  diffuse 
nucleus  internally  in  the  form  of  bars  or  rodlets  of  chromatin 
distributed  along  the  body.  In  some  forms  there  is  a  membrane  or 
crista  present  (fig.  53),  which  in  the  past  was  compared  with  the 
undulating  membrane  of  a  trypanosome,  but  the  membrane  of  a 
spirochaete  does  not  undulate.  Progression  is  very  rapid,  corkscrew- 
like and  undulatory  movements  occurring  simultaneously. 

The  genus  Spirochceta  was  founded  by 
Ehrenberg  in  1833  for  an  organism  which 
he  discovered  in  stagnant  water  in  Berlin. 
Ehrenberg  named  the  organism  Spirochceta 
plicatilis.  According  to  Zuelzer  (191 2)  S. 
plicatilis  does  not  possess  a  membrane  or 
crista,  but  an  axial  filament.  S.  gigantea  has 
been  described  by  Warming  from  sea- water. 

Spirochaetes  occur  in  the  crystalline  style 
and  digestive  tract  of  many  bivalve  molluscs. 
The  first  molluscan  spirochaete  to  be  studied 
was  that  of  the  oyster,  named  by  Certes  (1882) 
"  Trypanosoma  "  balhianii  (fig.  53).  Similar 
spirochaetes,  probably  belonging  to  the  same 
species,  occur  in  various  species  of  Tapes  and 
in  Pectcn  (the  scallop).  S.  halbianii  has  rounded 
ends  (fig.  53).  Other  spirochaetes  occur  in  fresh- 
water mussels  {Anodonta  spp).  .S.  anodonice, 
studied  by  Keysselitz  (1906)  and  by  Fantham 
(1907),  has  pointed  ends.  Gross  (191 1)  sug- 
gested the  generic  name  Cristispira  for  mol- 
luscan spirochaetes,  because  they  possess  a 
well-marked  membrane  or  "crista,"  which 
appears  to  be  absent  from  S.  plicatilis,  accord- 
ing to  Zuelzer's  researches. 

Schaudinn  in  1905  founded  the  genus 
Treponema  for  the  parasite  of  syphilis  {T. 
pallidum),  discovered  by  him  and  by  Hoffmann. 
According  to  Schaudinn  the  Treponemata  have  no  membrane  or 
crista.  The  pathogenic  agent  of  yaws  or  framboesia,  discovered  by 
Castellani,  is  also  placed  in  the  genus  Treponema,  as  T.  pertenue. 

There  remain  the  blood  spirochaetes.  It  is  somewhat  disputed  as 
to  whether  these  organisms  possess  a  membrane.  The  present  writer 
considers  that  they  have  a  slight  membrane  or  crista.     The  name  of 


Fig.  53. — SpirocJmta  bal- 
hianii. a,  basal  granule  or 
polar  cap.  /^,  chromatin  rod- 
lets,  r,  membrane  ("'crista"). 
d,  myonemes  in  membrane. 
■{After  Fantham  and  Porter.) 


THE  spirochj5:tes  115 

tlie  genus  in  which  to  place  the  blood-inhabiting  forms  is  somewhat 
uncertain  and  disputed.  Various  generic  names  given  to  them  are 
Spirochceta,  Treponema,  Spiroschaudinriia  (Sambon)  and  Borrelia 
(Swellengrebel).  Included  in  this  division  are  the  causal  agents  of 
relapsing  or  recurrent  fever.  These  Protists  will  be  named,  for 
description,  Spirochaetes  without  prejudice  as  to  the  ultimate  correct 
generic  name. 

It  is  sometimes  made  a  matter  of  argument  as  to  whether  the 
spirochcctes  are  Protozoa  or  Bacteria.  Such  arguments  are  somewhat 
unprofitable.  Morphologically  the  spirochaetes  are  like  the  Bacteria 
in  possessing  a  diffuse  nucleus.  They  differ  from  Spirilhim,  an 
undoubted  bacterial  genus,  in  being  flexible  and  not  possessing 
flagella.  Molluscan  spirochaetes,  however,  may  appear  to  have 
flagella  if  their  membrane  becomes  frayed  or  ruptured,  when  the 
myonemes  therein  (fig.  53),  becoming  separated,  form  apparent 
threads  or  flagella  (Fantham,   1907-08).^ 

Again,  the  mode  of  division  of  spirochaetes  has  been  used  as  a 
criterion  of  their  bacterial  or  protozoal  affinity.  They  have  been 
stated  to  divide  transversely,  longitudinally,  and  by  *'  incurvation," 
or  bending  on  themselves  in  the  form  of  a  U,  "a  form  of  transverse 
fission."  The  present  writer  believes  that  they  divide  both  transversely 
and  longitudinally,  and  that  there  is  a  periodicity  in  their  mode  of 
division  at  first  longitudinal  (when  there  are  few  spirochaetes  in,  say, 
the  blood)  and  then  transversely  (w^hen  spirochaetes  are  numerous  in 
the  blood).^  Some  authors  consider  that  longitudinal  division  is  ex- 
plained  by  :"  incurvation." 

The  spirochaetes  of  relapsing  fever  show  a  remarkable  periodic 
increase  and  decrease  in  numbers  in  the  blood.  They  are  transmitted 
bv  ticks  or  by  lice.  They  react  to  drugs  {e.g.,  salvarsan  or  ^'  606  ") 
rather  like  trypanosomes,  and — like  Protozoa,  but  unlike  Bacteria — 
they  are  cultivated  with  difficulty.  These  and  other  criteria  have  been 
used  to  endeavour  to  determine  whether  they  are  Protozoa  or  Bacteria. 
The  present  writer  believes  that  they  are  intermediate  in  character, 
showing  morphological  affinities  with  the  Bacteria  and  physiological 
and  therapeutical  affinities  with  the  Protozoa.  The  group  Spiro- 
chaetacea,  as  an  appendix  to  the  Protozoa,  has  been  created  for  them 
by  the  present  writer  (Jan.,  1908).  Others  have  placed  them  in  the 
Spirochaetoidea  of  the  Bacteria  or  with  the  Spirillacea.  Doflein  (1909) 
called  them  Proflagellata.  Further  discussion  is  unnecessary,  as  they 
are  undoubtedly  Protista  (see  p.  29). 

There  is  no  true  conjugation,  sex  or  encystment  in  spirochaetes, 
but  morphological  variation  may  occur.^     They  may  agglomerate. 

'  Qtiart.  Journ.  Microsc.  Set.,  lii,  p.  I.  ^  Froc.  Roy.  Soc,  B,  Ixxxi,  p.  500. 

3  Fantham,  Parasitology,  ii,  p.  392. 


Il6  THE   ANIMAL    PARASITES   OF    MAN 

The  Spirochaetes  form  an  interesting  chapter  in  the  evokition  of 
parasites.  There  are  free  Hving  forms,  parasitic  forms  in  the  guts 
of  both  vertebrates  and  invertebrates,  and  blood-inhabiting  forms. 
These  probably  represent  the  order  of  evolution  of  parasitism.  The 
blood-inhabiting  forms  are  pathogenic  to  warm-blooded  hosts. 

We  must  now  consider  the  blood  Spirochaetes  and  the  Treponemata 
(organisms  of  syphilis  and  of  yaws). 

THE  SPIROCHETES  OF  THE  BLOOD. 

There   are   at    least    two    important    human    parasites    included 
hereunder  ; — 
(a)  Spirochceta  recurrentls  (^S.  obermeieri),  (b)  Spirochcvta  diittoni. 
More  is  known  of  the  life-cycle  of  Spirochceta  dnttonij  and  it  will 
be  convenient  to  consider  that  first. 

Spirochaeta  duttoni,  Novy  and  Knapp,  1906. 

The  specific  name  duttoni  was  also  given,  independently,  to  this  parasite  in  1906 
by  BreinI  and  Kinghorn. 

S.  duttoni  is  the  pathogenic  agent  of  African  tick  fever  in  man, 
prevalent  in  the  Congo  State  and  other  parts  of  Africa.  The  full- 
grown  organism  is  about  16  //,  to  24  /i,  long,  and  has  pointed  ends.  It 
is  0*25  yu,  to  0*5  yLt  broad.  P.  H.  Ross  and  Nabarro  were  among  the 
earliest  to  see  a  spirochaete  in  the  blood  of  patients  in  Uganda.  It  is 
transmitted  by  the  tick,  Ornithodorus  moxibata. 

In  the  blood  of  the  patient  some  of  the  spirochaetes  may 
show,  after  staining,  lighter  and  darker  portions  (chromatin  dots)  and 
evidence  of  the  possession  of  a  very  narrow  membrane  (fig.  54). 
The  mode  of  division  has  already  been  discussed.  Periodicity  in 
the  direction  of  division  was  first  described  by  Fantham  and  Porter,^ 
(1909).  Just  before  the  crisis  in  African  tick  fever,  BreinI  has  stated 
that  S.  duttoni  becomes  thinner  in  the  spleen  and  bone-marrow 
and  rolls  up  into  skein-like  forms,  which  are  surrounded  by  a  thin 
*^cyst"  wall  (probably  the  periplast).  Such  occur  in  apyrexial 
periods.  Inside  the  cyst  the  spirochaete  breaks  up  into  granules. 
Balfour  and  Sambon  have  described  somewhat  similar  rolled  up 
forms,  breaking  into  granules,  inside  the  red  blood  cells  of  Sudanese 
fowls  in  the  case  of  S.  granulosa  (possibly  only  a  variety  of  S.  gal- 
linarum).     The  intracorpuscular  stage  is  not  definitely  established. 

The  granule  phase,  however,  is  an  essential  one  in  the  invertebrate 
transmitter  (fig.  54c).  In  1905,^  Button  and  Todd  proved  experimen- 
tally that  0.  nioubata  transmitted  S.  duttoni.     They  fed  ticks,  obtained 

*  Proc.  Roy,  Soc,  B,  Ixxxi,  p.  500. 

-  Liverpool  Sch.  Trop.  Med.,  Metnoir  xsn'.  Lancet,  Nov.  30,  1907,  p.  1523. 


SPIROCH^TA   DUTTONl 


117 


from  Congo  native  huts  in  which  infected  persons  hved,  on  monkeys 
and  the  latter  became  infected.  Button  and  Todd  also  found  the 
offspring  of  infected  ticks  to  be  capable  of  transmitting  the  infection 
to  experimental  animals.  They  concluded  that  0.  mouhata  was  a 
true  intermediate  host. 

A  little  later  in  1905,  Koch  stated  that  spirochaetes  from  the  gut 
of  the  tick  penetrated  the  gut  wall  and  tissues  and  found  their 
way  into  the  eggs  in  the  ovary.  Koch  figured  tangled  masses  of 
spirochaetes  as  occurring  in  the  tick  eggs.  He  found  ticks  infective 
to  the  third  generation.  He  thought  that  the  infection  was  spread 
by  the  salivary  fluid  of  the  tick,  in  the  act  of  biting.  (This  is  now 
known  to  be  incorrect.)  Markham  Carter  (1907)  corroborated  Koch's 
work  on  the  spirochaetes  in  the 
tick  eggs,  and  they  have  been 
seen  since  by  Kleine  and  Eckard 

(1913)- 

Sir  William    Leishman,^  in 

1909-10,  found  that  at  ordinary 

temperatures  the  salivary  glands 

of  infected  ticks  (0.  rnoiibata) 

were  not  themselves   infective, 

and  that  the  infection  occurred 

by  way  of  the  ticks'  excretion. 

The  spirochaetes  (contained  in 

the  ticks'  excrement)  found  their 

way    into    the    vertebrate    host 

through   the  wound   made   by 

biting.      While    feeding,    ticks 

pass   large   quantities    of    clear 

fluid    from   the   coxal   glands  ; 

in    this    fluid    an    anticoagulin 

occurs.       Some  of  the  ticks  also  Fig.    54.  —  Spirochxta    duttoni.      a,    blood 

pass     thick,     white     Malpighian  form  showing  slight  membrane;   <^,   granules  or 

^            .            [lu    i-     •                               +  coccoid  bodies  clearly  formed  wiihin  the  organism; 

secretion,     that     is,     excrement,  ^^  beginning  of  extrusion  of  coccoid  bodies  in  the 

towards  the  end    of   the    feed.     t'ck.    (After  Fantham.) 
LeishmaU;  using   experimental 

monkeys,  showed  that  if  infected  ticks  were  interrupted  while 
feeding,  then  no  infection  resulted  in  the  monkeys.  If,  however,  the 
ticks  were  allowed  to  finish  their  feed,  and  the  Malpighian  secre- 
tions were  passed,  then  the  experimental  monkeys  became  infected. 
Fantham^  and  Hindle^  (191 1)»  independently,  have  repeated  the 
experiments  with  mice. 


•% 


'  Journ.  Roy.  Army  Med.  Corps,  xii,  p.  123  ;  Lancet  (1910),  clxxviii,  p.  ii. 

'2  Anncds  Trop.  Med.  and  Parasitol.,  v,  p.  479.  '  Parasitology,  iv,  p.  133. 


ii8 


THE   ANIMAL   PARASITES   OF   MAN 


Coxal 


Leishman's  methods  and  results  may  be  summarized  thus  : 
SaHne  emulsions  of  the  organs  of  infected  ticks  were  made,  after 
the  organs  had  been  most  carefully  dissected  out.  The  ticks  were 
first  kept  for  several  days  at  certain  constant  temperatures,  such  as 
24°  to  25°  C.  or  blood  heat,  37°  C.  The  saline  emulsions  of  the 
organs  were  inoculated,  separately,  into  experimental  animals,  and  the 
results  recorded  : — 

At  24°  C.  At  37°  C. 

Salivary  glands  Negative  ...  Positive 

Malpighian  tubules Positive  ...  Positive 

Gut  and  contents       Positive  ...  Positive 

Excrement      Positive  ...  Positive 

Genital  organs  Positive  ...  Positive 

fluid   is   usually   negative  ;    thick,  white    excrement  from 

Malpighian  tubes  is  posi- 
tive. 

When  the  ticks  were 
incubated  at  21°  to  24°  C. 
no  spirochaetes,  as  such, 
were  seen  in  the  organs, 
except  perhaps  in  the  gut, 
where  they  often  dis- 
appeared in  a  few  days. 
When  the  ticks  were  pre- 
viously incubated  at  35°  to 
37°  C.  for  two  to  three  days, 
spirochaetes,  as  such,  re- 
appear in  the  gut,  organs 
and  haemoccelic  fluid. 
The  infection  proceeds, 
from  the  salivary  gland,  but  from  the  infective  excrement, 
is,  from   the   thick,  white   material    voided    by   the   tick   while 


Fig.  55. — Spirochceta  duttoni  and  its  coccoid  bodies 
in  the  tick  {0.  vioubata). — Mononuclear  cells  of  the  tick 
{O.  moubata)  containing  [a)  Spirochete  breaking  up  into 
coccoid  bodies ;  {b)  Similar  tick-cell  containing  coccoid 
bodies  or  granules.  Such  mononuclear  cells  occur  in 
various  organs  of  ticks  and  in  developing  Malpighian 
tubules.     (Original.     From  preparations  by  Fantham.) 


not 
that 


feeding,  usually  towards  the  end  of  the  meal.  This  Malpighian 
excrement  passes  into  the  wound  caused  by  the  bite,  being  greatly 
aided  by  the  clear  and  more  limpid  coxa  fluid,  which  bathes  the  under 
surface  of  the  tick's  body,  and  mixes  with  and  carries  the  infective 
excrement  into  the  wound.     Ticks  remain  infective  for  a  long  time. 

The  spirochaetes  in  the  gut  of  infected  ticks  divide  by  a  process 
of  multiple  transverse  fission  into  granules,  which  are  composed  of 
chromatin  (fig.  54).  These  granules — sometimes  known  as  coccoid 
bodies — are  capable  of  multiplication.  Leishman  first  found  them  in 
clumps  inside  the  cells  of  the  Malpighian  tubules  (cf.  fig.  55). 

To  summarize,  when  spirochaetes  are  ingested  by  a  tick,  some  of 
them  pass  through  the  gut- wall  into  the  haemoccelic  (body)  fluid. 
They  then  bore  their  way  into  the  cells  of  various  organs  (fig.  55^7) 


SPIROCH^TA   GALLINARUM  II9 

and  break  up  into  coccoid  bodies.  In  this  manner  the  granules  find 
their  way  into  the  ovaries  and  ova,  thus  explaining  how  the  young 
ticks  are  born  infected.  Inoculation  of  these  chromatinic  granules 
usually  produces  infection.  Infective  granules  are  also  seen  in  the 
rudiments  of  the  Malpighian  tubules  of  embryo  ticks.  Bosanquet 
and  Fantham  (191 1),  independently,  have  shown  that  molluscan 
spirochaetes  also  break  up  into  similar  granules  or  coccoid  bodies. 
Gross  has  also  demonstrated  multiple  transverse  fission  in  molluscan 
forms.  Marchoux  and  Couvy  (19 13)  and  Wolbach  (1914)  consider 
the  granules  or  coccoid  bodies  to  be  degeneration  products.  This  is 
unlikely  (see  below). 

Schuberg  and  Manteufel  have  found  that  certain  0.  moubata, 
perhaps  30  per  cent,  of  the  specimens  of  a  given  neighbourhood,  may 
acquire  a  natural  active  immunity  against  infection  with  S.  duttoni. 

S.  duttoni,  or  a  closely  allied  form  (by  some  termed  S.  novyi),  occurs 
in  Colombia,  and  is  spread  by  the  tick  Ornithodorus  turicata.  In 
Panama  a  similar  spirochaete  is  probably  spread  by  0.  talaje. 

Spirochseta   gallinarum,  Stephens  and  Christophers,   1905 
(=  Spirochaeta  marchouxi,  Nuttall,  1905). 

This  Spirochaete,  which  occurs  in  fowls  and  is  pathogenic,  is 
transmitted  by  the  tick  Argas  persicus.  It  is  about  10  fi  to  20  fj,  long. 
There  is  a  pathogenic  spirochaete  known  to  occur  in  geese,  named  by 
Sakharoff  (1891)  S.  anserina,  and  found  in  Caucasia.  This  may  be 
the  same  as  S.  gallinanim,  in  which  case  the  name  S.  anserina  will 
have  priority.  These  organisms  cause  fever,  diarrhoea,  anaemia  and 
death.  The  life  history  of  the  avian  pathogenic  spirochaetes  has  been 
studied  by  Balfour,  by  Hindle^  and  by  Fantham.^  It  is  essentially 
similar  to  that  of  S.  duttoni. 

Marchoux  and  Couvy^  (1913)  consider  that  the  '^fragmentation  of 
the  chromatin  "  in  spirochaetes  is  a  process  of  degeneration.  Working 
with  A.  persicus  and  S.  gallinarum,  they  state  that  a  large  number  of 
the  spirochaetes  ingested  by  the  Argas  almost  immediately  pass 
through  the  wall  of  the  alimentary  canal  and  appear  in  the  haemo- 
ccelic  fluid.  Marchoux  and  Couvy  consider  that  Leishman's  granules 
may  be  found  in  the  Malpighian  tubules  of  various  Arachnids.  They 
found  spirochaetes  in  the  cephalic  glands  of  infected  Argas.  They 
consider  that  spirochaetes  remain  as  wavy  spirochaetes  within  the  tick, 
if  they  are  to  be  infective,  though  the  spirochaetes  may  become  so  thin 
as  to  be  invisible  !  The  latter  argument  is  obviously  weak,  and  it 
was  never  asserted  that  all  granules  in  the  Malpighian  tubules  of 
infected   ticks  were  derived   from   spirochaetes.      With  dark-ground 

'  Parasitology,  iv,  p.  463.  ^  Annals  Trop.  Med.  and  Parasitol.  {1911),  v,  p.  479. 

^  Annales  Inst.  Pastenr,  xxvii,  pp.  450,  620. 


120  THE   ANIMAL   PARASITES   OP^   MAN 

illumination  small,  refractile  spirochaetal  granules  may  be  seen  to  grow 
into  spirochaetes.  The  granule  phase  of  spirochaetes  has  recently 
been  discussed  by  Fantham^  (1914)- 

Spirochaeta  recurrentis,  Lebert,  1874. 

Syn. :  Spirochceta  obermeieri^  Cohn,  1875. 

This  organism  was  discovered  by  Obermeier  (1873)  in  cases  of 
relapsing  fever  in  Berlin.  Short  forms  7 /x  to  9 /^  long,  and  longer 
(probably  adult)  forms,  16 /^  to  19//-,  are  found  in  the  blood.  The 
width  is  0-25  /x.     Parasites  12  yu,  or  13  /a  long  are  often  observed. 

The  spirochaete  is  found  in  the  blood  during  febrile  attacks  and 
relapses,  but  not  during  intervening  periods.  It  can  be  inoculated 
into  monkeys,  rats  and  mice.  It  can  live  in  the  bed-bug,  Cimex 
lectnlarms,  and  Nuttall  has  succeeded  in  transmitting  S.  recurrentis 
from  mouse  to  mouse  by  the  bites  of  the  same  bug.  The  French 
investigators  Sergent  and  Foley  (1908-9)  in  Algeria,  and  NicoUe, 
Blaizot  and  Conseil  (191 2)  in  Tunis,  have  shown  experimentally  that 
S.  recurrentis  (var.  berhera)  is  transmitted  by  lice.  The  latter  workers 
also  demonstrated  the  method  of  infection  that  commonly  occurs, 
namely,  by  the  scratching  of  the  skin  and  crushing  of  lice  containing 
spirochaetes  on  the  excoriated  surface  of  the  body. 

Lice  as  transmitting  agents  for  relapsing  fever  were  indicated  by  Mackie^  in 
1907.  An  epidemic  among  Indian  school  children  furnished  the  materials.^  It  was 
noted  that  out  of  170  boys,  137  were  infected,  and  the  boys  were  very  verminous. 
Among  the  girls,  35  out  of  114  suffered,  and  few  lice  were  found  on  them.  Twenty- 
four  per  cent,  of  the  lice  taken  from  the  boys  contained  spirochetes  as  compared 
with  3  per  cent,  of  those  from  the  girls.  As  the  epidemic  died  out  among  the  boys, 
the  lice  also  became  fewer>  and  an  increase  in  the  number  of  cases  among  the  girls 
coincided  with  an  increase  in  the  number  of  lice.  Spirochaetes  were  found  in  the 
gut,  Malpighian  tubules  and  genital  organs  of  the  lice.  Mackie  thought  that  infec- 
tion of  the  patients  was  brought  about  by  the  regurgitation  of  the  spirochaetes  when 
the  lice  fed,  but  proof  of  this  was  lacking. 

In  191 2,  Nicolle,  Blaizot  and  Conseil,*  working  in  Tunis  and 
using  chiefly  an  Algerian  strain  of  relapsing  fever  spirochaetes  (some- 
times called  S.  berhera) j  showed  by  direct  experiments  that  infection 
by  means  of  the  bites  of  Pediculus  vestinienti  and  P.  capitis  was 
untenable.  As  many  as  4,707  infected  lice  were  fed  on  one  man,  and 
6,515  on  another  occasion  were  allowed  to  bite  a  man  after  they 
had  fed  on  a  monkey  heavily  infected  with  spirochaetes,  yet  no 
infection  of  the  man  followed.  Examination  of  the  lice  showed 
that  the  spirochaetes  left  the  gut  soon  after  they  were  ingested,  and 

'  Amials  Trop.  Med.  and  ParasitoL,  viii,  p.  471. 

"^  Brit.  Med.  Journ.,  Dec.  14,  1907,  p.  1706. 

'  See  also  Nuttall,  Herter  Lecture  on  Spirochsetosis,  Parasitology,  v,  p.  269. 

*  C.R.  Acad.  Sci.,    cliv,  p.  1636  ;  civ,  p.  481. 


SPIROCH^TA   RECURRENTJS  121 

passed  into  the  body  cavity,  which  swarmed  with  spirochaetes.  The 
contents  of  the  ah'mentary  tract  and  the  faeces  of  the  Hce  ahke  were 
iininfective.  The  spirochaetes  did  not  reappear  in  the  gut  till  eight 
days  after  an  infective  feed,  but  some  persisted  as  late  as  the  nineteenth 
day  when  kept  at  28°  C. 

It  was  noted  that  the  irritation  due  to  the  lice  caused  scratching, 
and  that  thereby  lice  became  crushed  on  to  the  skin.  An  emulsion 
was  made  of  two  infected  lice  and  rubbed  on  to  the  slightly  excori- 
ated skin  of  one  of  the  above  workers.  Infection  followed  five  days 
later.  A  drop  of  emulsion  placed  on  the  conjunctiva  of  the  human 
eye  produced  spirochaetosis  after  an  incubation  of  seven  days.  The 
body  contents  of  such  lice,  then,  produce  infection  when  they  reach 
the  blood  by  any  excoriated  or  penetrable  surface.  The  stages  leading 
lip  to  infection  in  nature  briefly  are  :  The  irritation  due  to  the  louse 
bites  causes  scratching,  and  the  lice  are  crushed  on  to  the  skin.  The 
slight  abrasion  is  quite  sufficient  to  permit  the  entry  of  the  parasite. 
The  louse  bite  alone  is  harmless.  Infection  by  way  of  the  eye  is 
quite  probable  in  Africa,  remembering  the  constant  trouble  due  to 
sand,  dust,  insects,  etc.,  resulting  in  frequent  touching  of  the  eyes. 

The  spirochaetes  occur  in  the  body  fluid  of  the  lice  and  can  pass 
in  it  to  the  adjacent  organs.  Thus  they  probably  find  their  way  into 
the  genital  organs,  and  into  the  eggs  of  the  lice.  Eggs  laid  twenty 
to  thirty  days  after  the  parent  became  infected  have  retained  the 
infection,  and  the  larvae  issuing  from  such  eggs  must  have  contained 
some  form  of  spirochaetes,  for  an  emulsion  of  either  the  eggs  or 
the  larvae  produced  spirochaetosis  when  inoculated  into  monkeys. 
Further  details  regarding  the  spirochaetosis  in  the  eggs  of  the  lice  and 
in  the  larvae  are  needed.  Hereditary  infection,  however,  has  been 
demonstrated,  but  is  not  very  common.  Sergent  and  Foley  (19 14) 
state  that  the  spirochaete  possesses  a  very  small  and  virulent  form 
which  it  assumes  during  apyrexial  periods  in  man  and  during  a 
period  following  an  infecting  meal  in  the  louse.  Nicolle  and  Blanc 
{1914)  find  that  the  organisms  are  infective  in  the  louse  just  before 
they  reappear  as  spirochaetes.  Nicolle  and  Blaizot  found  that  female 
lice  were  more  susceptible  to  spirochaetes  than  males,  four  times 
as  many  females  as  males  being  infected. 

Tictin  (1897)  found  S.  recurrentis  in  bugs  recently  fed  on  patients,  and  infected  a 
monkey  with  the  fluids  of  crushed  bugs.  Karlinski  (1902)  found  the  spirochaetes 
in  bed-bugs  in  infected  houses.  There  is  some  other  evidence  to  show  that  bugs 
may  transmit  the  spirochaete  in  Nature.  Further  researches  are  needed  regarding 
the  relationship  of  bed-bugs  and  human  spirochaetosis. 

Multiplication  of  S.  recurrentis  is  by  longitudinal  and  transverse 
division  (including  so-called  '^incurvation"),  and  the  organism  forms 
small,  ovoid  bodies  (''  coccoid  "  bodies)  in  the  same  way  as  S.  diittoni. 


122  THE   ANIMAL   PARASITES   OF   MAN 

S.  recurrentis  is  the  cause  of  European  relapsing  fever,  and  a 
number  of  possible  varieties  of  it  are  associated  with  relapsing  fevers 
in  other  parts  of  the  world.  Such  spirochaetes  only  differ  by  bio- 
logical reactions,  such  as  acquired  immunity  tests.     They  include  : — 

5.  rossii^  the  agent  of  East  African  relapsing  fever  ;  S.  novyiy  the 
agent  of  North  American  relapsing  fever;  S.  carteri,  the  agent  of 
Indian  relapsing  fever;  S.  herhera,  the  agent  of  North  African  and 
Egyptian  relapsing  fever. 

Other  Human  Spirochetes  are: — 

S.  schaiidinni.  This  organism,  according  to  Prowazek,  is  the 
agent  of  ulcus  tropicum.     It  varies  in  length  from  lo/i  to  20 /x. 

S.  ahoriginalis  has  been  found  in  cases  of  granuloma  inguinale 
in  British  New  Guinea  and  Western  Australia.  It  also  occurs  in 
dogs,  and  may  not  be  truly  parasitic. 

S.  vincenti.  This  spirochaete  is  12  /a  to  25  /i,  in  length,  tapers  at 
both  ends  and  has  few  coils.  It  has  been  associated  with  angina 
vincenti.     It  often  occurs  in  company  with  fusiform  bacilli. 

S.  hronchialis,  found  by  Castellani  in  1907  in  cases  of  bronchitis 
in  Ceylon.  The  parasites  are  delicate,  but  show  morphological 
variation.  This  organism  is  important  and  has  since  been  found 
in  the  West  Indies,  India,  Philippine  Islands  and  various  parts  of 
Africa,  such  as  the  Anglo-Egyptian  Sudan,  Uganda  and  West  Africa. 
It  has  recently  been  the  subject  of  research  by  Chalmers  and 
O'Farrell,  Taylor,  and  Fantham. 

S.  phagedenis  was  found  by  Noguchi  in  a  ten  days  old  ulcerated 
swelling  of  the  labium.  The  organism  shows  much  variation  in  size, 
being  4/x  to  30  //,  in  length. 

S*  refringens  (Schaudinn,  1905)  occurs  in  association  witli 
Treponema  pallidum  in  syphilitic  lesions,  but  is  non-pathogenic.  It 
is  20  /i,  to  35  /A  long  and  0*5  jju  to  075  fi  broad,  being  larger  than 
r.  pallidum  and  more  easily  stained. 

Various  spirochaetes  have  also  been  notified  in  vomits,  chiefly  in 
Australia  ;  others  from  the  human  intestinal  tract,  e.g.,  S.  eurygyrata  ; 
S.  stenogyrata  (Werner) ;  S.  hachaizce  (Kowalski),  in  cholera  motions; 
S,  buccalis  (Cohn,  1875)  and  S.  detitiiim  occurring  in  the  human 
mouth  and  in  carious  teeth  (S.  dentium,  Koch,  1877,  being  the 
smaller) ;  S.  acuminata  and  S.  obtusa  found  by  Castellani  in  open 
sores  in  cases  of  yaws. 

Animal  spirochaetes  of  economic  importance  include  : — 

S.  anserinay  highly  pathogenic  to  geese. 

S.  gallinariim    ( =  S.    marchouxi)   in  fowls.     (See  p.  119.) 

S.  theileri  in  cattle  and  S.  ovina  in  sheep  also  occur  in  Africa ; 
their  pathogenicity  is  not  clear. 

S.  laverani  (  =  S.  miiris),  occurring  in  the  blood  of  and  patho- 


CULTIVATION   OF   SPIROCHETES  1 23 

genie  to  mice,  is  probably  the  smallest  spirochsete  from  the  blood, 
being  only  3 /x  to  6  fi  long. 

Numerous  spirochaetes  have  been  recorded  from  the  guts  of 
various  mammals,  birds,  fishes,  amphibia  and  insects. 

Cultivation  of  Spirochetes. — Cultures  of  spirochaetes  have 
been  made  with  little  success  or  with  great  difficulty  until  compara- 
tively recently,  when  Noguchi  (191 2)  devised  a  means  whereby  he 
has  cultivated  most  of  the  pathogenic  spirochaetes  as  well  as  some 
Treponemata. 

Noguchi  has  now  cultivated  S.  duttoni,  S.  recurrentis,  S.  rossii, 
S.  novyi  and  S.  gallinariun  from  the  blood ;  S.  phagedenis^  from  human 
phagedaenic  lesions  ;  S.  refringens^  and  spirochaetes  from  the  teeth. 

His  method  is  as  follows  : — 

A  piece  of  fresh,  sterile  tissue,  usually  rabbit  kidney,  is  placed  in  a 
sterile  test-tube.  A  few  drops  of  citrated  blood  from  the  heart  of  an 
infected  animal,  ^.^.,  rat  or  mouse,  is  added,  and  about  15  c.c.  of  sterile 
ascitic  or  hydrocoele  fluid  is  poured  quickly  into  the  tube.  Some  of 
the  tubes  are  covered  with  a  layer  of  sterile  paraffin  oil,  others  are 
left  uncovered.  The  tubes  are  incubated  at  37°  C.  The  best  results 
are  obtained  if  the  blood  is  taken  from  an  animal  forty-eight  to  seventy- 
two  hours  after  it  has  been  inoculated,  that  is,  before  the  spirochaetes 
reach  their  maximum  multiplicative  period  in  the  blood.  The  presence 
of  some  oxygen  seems  indispensable  for  these  blood  spirochaetes,  and 
they  fail  to  develop  in  vacuo  or  in  an  atmosphere  of  hydrogen. 

For  subcultures,  0*5  c.c.  of  a  culture  is  added  to  the  medium 
instead  of  citrated  blood,  and  it  is  useful  to  add  a  little  fresh,  normal 
blood,  either  human  or  from  an  animal,  such  as  a  rat. 

Noguchi  found  that  the  events  in  cultures  were  :^ 

S.  duttoni,^  maximum  multiplication  on  the  eighth  to  ninth  day  ; 
disintegration  beginning  on  the  tenth  day,  spirochaetes  disappeared 
after  about  the  fifteenth  day.  No  diminution  of  virulence  was  found 
at  the  ninth  day. 

S.  rossii  (  =  S.  kochi).^  Maximum  development  on  the  ninth  day, 
after  which  the  virulence  diminishes.  The  incubation  period  is  also 
prolonged. 

S.  recurrentis^  (=S.  obermeieri).  Maximum  growth  on  the  seventh  day. 

S.  novyi. ^ — Maximum  development  on  the  seventh  day.  It  is 
more  difficult  to  grow  than  the  preceding  forms. 

All  the  above  spirochaetes  showed  undoubted  longitudinal  division 
and  transverse  division  was  observed  in  part. 

S.  gallinarum  ^  can  be  cultivated  as  above,  but  transverse  division 

^  Jotirn.  Exptl.  Med.,  xvi,  p.  261.  ^  Journ.  Exptl.  Med.,  xv,  p.  466. 

^  Journ.  Exptl.  Med.,  xvi,  p.  202.  ■•  Ibid.,  p.  205. 

^  Ibid.,  p.  205.  «  Ibid.,  p.  20S.  "  Ibid.,  p.  620. 


124 


THE   ANIMAL   PARASITES   OF   MAN 


was  usual  here.     Maximum  growth  occurred  in  the  culture  about  the 
fifth  day. 

Treponemata. 

The  genus  Treponciim  (Schaudinn,  1905),  includes  minute,  thread- 
like organisms,  with  spirally  coiled  bodies,  the  spirals  being  preformed 
or  fixed.  No  membrane  or  crista  is  present,  according  to  Schaudinn, 
though  a  slight  one  is  said  by  Blanchard  to  be  present  in  the  organism 
of  yaws.  The  ends  of  the  organisms  are  tapering  and  pointed. 
Multiplication  is  by  longitudinal  and  transverse  division.  The  most 
important  members  of  the  genus  are  T.  pallidum,  the  agent  of  syphilis, 
and  T.  perte/iiie,  which  is  responsible  for  framboesia  or  yaws. 


Treponema  pallidum,  Schaudinn,  1905. 

Syn.  :    Spirochcrta  pallida. 

Treponema  pallidum  was  first  described  by  Schaudinn  and   Hoff- 
mann in  1905  under  the  name  of  Spirochaita  pallida.     It  has  also  been 

described  under  the  names 
oiSpirouema  pallida,  Micro- 
sp  iron  em  a  pallida  and 
Trypanosoma  hits.  Siegel 
in  1905  described  an  organ- 
ism which  he  called  Cyto- 
rhyctes  Inis  and  considered 
to  be  the  agent  of  syphilis. 
Schaudinn  reinvestigated 
Siegel's  work  and  found 
T.  pallidum,  which  he  con- 
sidered to  be  the  causal 
agent  of  the  disease,  and 
pronounced  against  Cyto- 
rhyctes  Inis.  It  is  probable 
now  that  both  workers 
were  correct,  for  Balfour 
(191 1)  has  seen  the  emis- 
sion of  minute  granules 
or  "  coccoid  "  bodies  from  T,  pallidum  and  these  granules  probably 
correspond  to  the  C.  luis  of  Siegel.  Recently  E.  H.  Ross,  having 
observed  a  spirochaete  stage  in  the  development  of  Kurloff  bodies, 
thinks  that  T.  pallidum  is  a  stage  in  the  life-history  of  a  Lympho- 
cytozoon.  MacDonagh  has  also  described  a  complicated  and  some- 
what similar  cycle,  but  these  observations  require  further  study  and 
confirmation. 

T.  pallidum  varies  from  4/x  to   10 fjL  in  length,  its  average  length 
being  y  fM,  while  its  w4dth  is  usually  about  0*25  /n.     Longer  individuals 


Fig.  56. —  Trepo7teina  pallidum.     (After  Bell,  from 
Castellani  and  Chalmers.) 


TREPONEMA    PALLIDUM 


125 


of  i6yLt  to  20 /x  have  been  recorded.  The  body  has  from  eight  to  ten 
spiral  turns  and  forms  a  tapering  process  at  each  end  (fig.  56).  The 
organism  is  most  difficult  to  stain,  and  its  internal  structure  is  Uttle 
known.  It  is  possibly  like  that  of  Spirochceta  diittoiil  or  S.  halhianiiy 
as  the  ''  granule  shedding  "  observed  by  Balfour  is  strongly  suggestive 
of  the  formation  of  resistant  bodies  by  those  spirochaetes.     Hoffmann 

(19 1 2)  has  seen  the  formation  of  spores  in  T.  pallidum. 
The  Treponemata  occur  in  the  primary 

and  secondary  sores,  but  are  difficult  to 
find  in  the  tertiary  eruptions  of  syphilis. 
Noguchi    and    Moore    (1913)   and    Mott' 

(19 1 3)  have  demonstrated  T.  pallidum  in 
the  brain  in  cases  of  general  paralysis  of 
the  insane.  Marie  and  Levaditi  (19 14), 
however,  consider  that  the  treponeme 
found  in  the  brain  in  such  cases  is 
different  from  T.  pallidum. 

Cultivation  of  T.  pallidum. — This  has 
been  accomplished  successfully  by  No- 
guchi,^ using  a  modification  of  his 
method  for  spirochaete  cultivation,  for 
T.  pallidum  is  much  more  difficult  to 
grow  than  spirochaetes,  being  a  strict 
anaerobe. 

The  apparatus  consists  of  two  glass 
tubes,  the  upper  being  connected  to  the 
lower  by  a  narrower  tube  passing  through 
a  rubber  cork  (fig.  57).  Both  tubes  are 
carefully  sterilized. 

A  piece  of  fresh,  sterile  rabbit's  kidney 
is  placed  in  the  lower  tube,  which  \s 
filled  with  ascitic  fluid,  or  ascitic  fluid 
and  bouillon  mixture.  The  tube  is  in- 
oculated with  syphilitic  material  and 
corked  by  inserting  the  upper  tube.  In  the  bottom  of  the  upper  tube 
a  piece  of  sterile  rabbit's  kidney  is  placed  and  syphilitic  material 
poured  over  it.  A  mixture  of  one  part  ascitic  fluid  and  two  parts  of 
slightly  alkaline  agar  is  then  poured  over  the  tissue  and  allowed  to 
solidify.  When  solid,  a  layer  of  sterile  paraffin  oil  is  poured  on  top 
of  it,  and  the  top  plugged  with  cotton  wool  (fig.  57).  The  whole  is  then 
incubated  at  37°  C.  for  two  or  three  weeks.  The  tissue  removes  traces 
of  oxygen  from   the   lower   levels   of  the  medium  and  also   probably 

'  Brit.  Med.  [ourn.^  Nov.  15,  1913.  p.  1,271. 
2  /ourn.  Exptl.  Med.,  xv,  p.  90;  xvi,  p.  211. 


7iraf/tn 
V    oil 


Solid 
^•medium 


^■Tissue 


nuid 

r-  niQdium 


|«-Txs6ue 


Fig.  57. — Diagram  of  apparatus 
for  cultivation  of  Treponema  palli- 
dum by  Noguchi's  method.  (After 
Noguchi.) 


126  THE   AxMIMAL   PARASITES   OF   MAN 

provides  a  special  form  of  nourishment.  At  first  T.  pallidum  grows  in 
the  soHd  medium,  and  then  when  the  cultural  conditions  in  the  lower 
fluid  portion  become  favourable,  the  organisms  migrate  thither  and 
multiply  abundantly.  At  first  the  culture  is  impure,  but  after  several 
transferences  a  pure  culture  is  obtained  readily. 

The  syphilitic  material  for  culture  is  prepared  by  cutting  off 
pieces  of  tissue  from  the  lesions,  washing  in  sterile  salt  solution 
containing  i  per  cent,  sodium  citrate,  and  then  emulsifying  the 
tissue  in  a  mortar  with  sodium  citrate. 

Good  cultures  shaw  rapid  multiplication,  w^hich  is  invariably 
by  longitudinal  division. 

In  his  various  cultivation  experiments  Noguchi^  found  morpho- 
logical and  pathogenic  variations  in  T.  pallidiun.  Three  forms  of  the 
organism  were  found,  namely,  thicker,  average  and  thinner  types. 
The  lesions  caused  in  the  testicle  of  the  rabbit  differ  according  to 
the  variety  inoculated,  but  more   work   is  necessary  on   the  subject. 

Noguchi^  has  cultivated  a  separate  organism,  T.  caUigyrum, 
from  tlie  surface  of  human  genital  or  anal  lesions,  either  syphilitic 
or  non-syphilitic.  It  is  apparently  non-pathogenic,  and  is  6  //,  to 
14  /x  long. 

Hata(i9i3)^  has  modified  the  Noguchi  technique  for  the  cultivation 
of  spirochaetes  and  treponemes,  with  a  view  to  simplification  and 
convenience.  Hata  substitutes  normal  horse  serum  for  ascitic  fluid 
and  the  "  buffy  coat "  of  the  clot  of  horse  blood  in  place  of  the  small 
pieces  of  rabbit's  kidney.  It  is  unnecessary  to  place  sterile  paraffin  on 
the  surface  of  the  medium. 

The  horse  serum  is  mixed  with  twdce  its  volume  of  physiological 
saline  solution.  The  mixture  is  placed  in  tubes  which  are  heated  on  a 
water-bath  at  58°  C,  the  temperature  being  raised  gradually  until  it 
reaches  70°  or  71°  C.  in  three  hours.  The  tubes  are  then  heated  at 
71°  C.  for  half  an  hour.  After  cooling,  the  contents  will  consist  of  an 
opaque  semi-coagulated  mass.  This  semi-coagulated  serum  and  saline 
mixture  may  be  substituted  for  Noguchi's  ascitic  fluid. 

The  buff  coagulum  is  cut  into  small  pieces,  about  i  c.c.  in  volumcx 
They  must  be  forced  with  a  sterile  glass  rod  to  the  bottom  of  the  semi- 
coagulated  serum  and  saline  mixture.  The  medium  is  inoculated  wath 
a  small  quantity  of  infected  blood  and  kept  at  37°  C.  In  the  case  of 
S.  recurrentis,  growth  of  spirochaetes  is  observed  on  the  second  day, 
reaching  a  maximum  in  five  to  seven  days.  The  growth  of  the 
organisms  proceeds  rather  more  slowly,  they  live  for  a  longer  period 
and  maintain  their  virulence  better  than  in  Noguchi's  medium. 


^  [ourn.  Exptl.  Afed.,  xv,  p.  201.  ^  /ourn.  Exptl.  Med.y  xvii,  p.  89. 

^  Centralbl.  f.  Bakt.,  Orig.,  Ixxii,  p.  107. 


TREPONEMA    PERTENUE  1 27 

Treponema  pertenue,  Caslellani,  1905. 

Syn. :  Spirochceta  pertenids ;  S.  pallidula^  Castellani,  19015. 

Castellani  discovered  the  organism  in  1905,  in  scrapings  of  yaws 
pustules.     He  first  described  it  under  the  name  of  Spirochceta  pertenuis. 

Treponema  pertenue  (fig.  58),  though  delicate  and  slender,  shows 
great  morphological  variation  both  in  length  and  thickness.  It 
may  be  short,  e.g.,  j /jl,  but  can  attain  18//,  to  20 /jl  in  length  and  may 
be  even  larger.  In  cultures  made  by  Noguchi,  thick,  medium  and 
thin  forms  were  found,  each  giving  rise  to  a  different  type  of  fram- 
boesial  lesion  when  inoculated  into  the  testicles  of  rabbits,  thus 
suggesting  the  possibility  of  the  occurrence  of  varieties  of  T.  pertenue. 

The    organism    is     difficult    to     stain,    but    occasionally    deeper 
staining  granules   are   found    along   its   body.     They  may  represent 
a    diffuse    nucleus.      Granule    formation 
similar  to  that  of    T.  pallidum  has   been 
observed  by  Ranken,  using  dark-ground 
illumination. 

Many  experiments  have  been  made 
with  a  view  to  establishing  the  identity 
of  the  organism  of  yaws  and  also  of 
differentiating  between  the  causative 
agents  of  yaws  and  syphilis.  Both 
monkeys  and  the  human  subject  have 
been  experimentally  inoculated  with  yaws 
material  and  have  developed  the  disease. 

In  an  early  experiment,  negroes  were 
inoculated  with  the  secretion  from  lesions 

.  All         r     ii  11  11  Fig.  58. —  freponema  pertenue. 

of     yaws.         All     of     them     developed    the       (After    Castellani  and  Chalmers.) 

disease,  nodules  appearing,  chiefly  at  the 

seat  of  inoculation,  in  from  twelve  to  twenty  days,  followed  by  the 

usual  eruption.     Similar  results  were  obtained  with  thirty-two  Chinese 

prisoners,  who   were   inoculated  with    yaws,  twenty-eight    becoming 

infected. 

A  naturally  infected  yaws  patient  when  inoculated  with  syphilis, 
contracted  that  infection,  thus  showing  that  yaws  does  not  confer 
immunity  to  syphilis.  This  has  also  been  observed  naturally,  when 
yaws  patients  have  contracted  syphilis. 

Experiments  with  monkeys  have  been  successfully  performed. 
The  incubation  period  varies  from  sixteen  to  ninety-two  days. 
Lesions  appear  first  at  the  seat  of  inoculation,  and  in  some 
monkeys  the  eruption  is  localized  to  this  spot,  though  the  infection  is 
general,  T.  pertenue  occurring  in  the  spleen,  lymphatics,  etc.  Monkeys 
inoculated  with  splenic  blood  of  a  yaws  patient,  and  also  sometimes 
with  blood  from  the  general  circulation,  have  become  infected. 


128  THE   ANIMAL   PARASITES   OF   MAN 

Castellani  and  others  have  shown  that  monkeys  successfully 
inoculated  with  syphilis  do  not  become  immune  to  yaws,  and  vice- 
versa. 

Craig  and  Ashburn,  using  the  monkey  Cynouiolgns  philippincnsis, 
found  these  animals  susceptible  to  yaws  but  not  to  syphilis. 

The  ulcerated  lesions  of  framboesia  are  rapidly  invaded  by 
numerous  bacteria  as  well  as  by  different  spirochaetes,  of  which 
Castellani  has  described  three  distinct  species.  One  is  identical 
with  Spirochceta  refringens,  Schaudinn,  the  other  two  are  thin  and 
delicate.  One,  S.  ohtiisay  has  blunt  ends ;  the  other  S.  acuminata,  has 
pointed  ends.     T.  pertenue  is  also  present. 

The  reasons  for  considering  T.  pertenue  to  be  the  specific  cause  of 
framboesia  are : — 

(i)  T.  pertenue  is  the  only  organism  present  in  non-ulcerated 
papules,  in  the  spleen  and  in  the  lymphatics  of  yaws  patients,  or 
of  monkeys  artificially  infected  with  the  disease.  By  no  method 
has  any  other  organism  been  obtained. 

(2)  Extract  of  framboesia  material,  free  from  all  organisms  other 
than  T.  pertenue,  reproduces  the  disease  if  inoculated. 

(3)  Extract  of  framboesia  material  deprived  by  filtration  of 
T.  pertenue  is  no  longer  infective  on  inoculation. 

The  method  of  infection  is  contaminative,  by  direct  contact. 
Women  in  Ceylon  are  frequently  infected  by  their  children.  Any 
slight  skin  abrasion  is  sufficient  to  admit  the  parasite.  In  some 
cases,  insects  may  carry  the  disease  from  person  to  person,  and  even 
in  hospitals,  when  dressings  are  removed,  it  has  been  noticed  that 
flies  greedily  suck  the  secretion  from  the  ulcers.  T.  pertenue  has 
been  recovered  from  flies  that  have  fed  on  yaws,  and  monkeys  have 
contracted  the  disease  when  flies  were  placed  and  retained  on  them 
for  a  short  time,  after  the  insects  had  fed  on  yaws  material. 

Cultivation. — T.  pertenue  has  been  cultivated  by  Noguchi,  who 
finds  three  types  of  parasites  in  his  cultures,  as  before  mentioned.  Its 
multiplication  is  by  longitudinal  division. 

Noguchi^  (191 2),  has  cultivated  species  of  Treponema  from  the 
human  mouth,  e.g.,  T.  ntacrodentium,  T.  niicrodeutluni  and  T.  mucosnni, 
the  latter  from  pyorrhea  alveolaris.  These  parasites  in  the  past  may 
have  been  confused  under  the  name  Spirochceta  dentium. 

Class  III.     SPOROZOA,  Leuckart,  1879. 
The  third  group   of   the   Protozoa    consists   entirely  of    parasitic 
organisms    forming    the    class    know^n    as   the  Sporozoa    or    spore- 
producing  animals.     The   members  of   this   class   are   characterized 

'  foiirn.  Exptl.  Med.,  xv,  p.  8l  ;  xvi,  p.  194. 


TELOSPORIDIA  129 

by  possessing  very  great  powers  of  multiplication,  coupled  with  a 
capacity  for  producing  forms  that  serve  for  the  transference  of  the 
organisms  to  other  hosts.  These  reproductive  bodies,  whether  for 
increase  of  numbers  within  one  host  or  for  transmission  to  another 
host,  are  called  spores.  But,  strictly,  the  term  spore  should  be  used 
only  in  the  latter  connection,  when  a  protective  or  resistant  coat 
known  as  a  sporocyst  envelops  the  body  of  the  spore. 

The  Sporozoa  are  widely  distributed,  occurring  in  various  tissues 
and  organs  of  Annelids,  Molluscs,  Arthropods,  and  Vertebrates. 
Their  food,  which  is  fluid,  is  absorbed  osmotically.  The  life-cycle 
of  a  Sporozoon  may  be  completed  within  one  host  or  may  be  dis- 
tributed between  two  different  hosts. 

The  Sporozoa  were  divided  by  Schaudinn  into  two  groups  or 
sub-classes,  called  (i)  the  Telosporidia,  and  (2)  the  Neosporidia. 

The  Telosporidia  are  Sporozoa  in  which  the  reproductive  phase 
of  the  parasites  is  distinct  from  the  growing  or  trophic  phase,  and 
follows  after  it.  The  Neosporidia  include  Sporozoa  in  which  growth 
and  spore-formation  go  on  simultaneously.  This  classification  is  not 
final,  for  certain  exceptions  and  difficulties  are  already  known  with 
regard  to  it.  It  is  possible  that  the  class  Sporozoa  is  not  a  natural 
entity,  but  should  be  replaced  by  two  classes  of  equal  rank,  corre- 
sponding in  most  respects  with  the  Telosporidia  and  Neosporidia. 

The  Telosporidia  comprise  the  Gregarinida,  the  Coccidiidea,  and 
the  Haemosporidia.  Doflein  combines  the  two  latter  orders  into  one 
known  as  the  Coccidiomorpha. 

The  Neosporidia  comprise  the  Myxosporidia,  the  Microsporldia, 
the  Actinomyxidia,  the  Sarcosporidia,  and  the  Haplosporidia. 
Doflein  combines  the  first  three  orders  into  one,  the  Cnidosporidia. 

Sub-Class.     TELOSPORIDIA,  Schaudinn. 

Sporozoa  in  which  the  reproductive  phases  follow  completion  of 
growth. 

Order.     Gregarinida,  Aime  Schneider  emend.  Doflein. 

Knowledge  of  the  Gregarinida  probably  goes  back  as  far  as  the  year  1684,  when 
Redi  observed  gregarines  in  the  crab,  Cancer  pagiirus.  Von  Cavolini  (1787)  found 
them  in  Cancer  depressus.  The  name  Gregarina  was  created  by  L.  Dufour  (1828), 
who  observed  masses  of  these  organisms  in  the  gut  of  insects  of  different  orders. 
Hammerschmidt  (1838)  and  von  Siebold  found  rich  infestations  in  insects,  while 
Dujardin  (1835)  and  Henle  described  various  genera  from  segmented  worms. 
Henle  (1835)  also  observed  cysts  containing  "navicellae"  in  the  sperm-sacs  of 
segmented  worms,  and  attention  was  drawn  to  his  researches  by  the  discovery  by 
von  Siebold  (1839)  of"  pseudonavkrellse"  in  the  gut  of  Sciara  nitidicollis.  Up  to  this 
time  many  workers  considered  the  gregarines  to  be  worms,  but  Kolliker  (1845) 
investigated  many  of  them  and  maintained  their  unicellular  nature,  while  Stein's 
work  (1848)  showed  the  interrelation  of  the  pseudonavicellas  and  the  gregarines. 
9 


IjO 


THE   ANIMAL   PARASITES   OF   MAN 


The  discovery  of  amoeboid  germs  in  the  pseudonavicellas  by  Lieberkiihn  (1855)  and 
the  demonstration  of  myonemes  further  aided  in  the  elucidation  of  their  true 
systematic  position.  The  entire  process  of  conjugation,  of  which  Du four  had  seen 
one  phase,  was  followed  by  Giard  under  the  microscope. 

From  1873  onwards  Aime  Schneider  made  important  additions  to  the  knowledge 
of  the  morphology,  life-history,  and  systematic  position  of  numerous  gregarines. 
Butschli  (1881)  and  L.  Leger  (1892)  also  contributed  much  work  on  the  subject.  The 
discoveries  of  Schaudinn  with  regard  to  the  life-cycle  of  Coccidia  gave  a  fresh 
stimulus  to  the  study  of  the  Gregarines,  whereby  the  life-cycles  of  numerous  forms 
and  the  phases  thereof  have  been  elucidated. 

Asexual  multiplication  is  not  common  among  the  Gregarines,  but  is  known  to 
occur  in  the  sub-order  Schizogregarinea,  formerly  known  as  the  Amoebosporidia. 

Although  the  Gregarinida  are  not  known  to  be  parasitic  in  man  or  other  verte- 
brates, they  are  of  great  interest,  inasmuch  as  they  are  among  the  earliest  known 
Sporozoa,  and  therefore  will  be  briefly  described  here. 

The  Gregarines  are  usually  elongate,  somewhat 
flattened  organisms  (figs.  59,  60),  whose  bodies  are 
enclosed  in  an  elastic  and  often  thick  cuticle.  The 
enclosed  living  substance  shows  a  separation  into 
ectoplasm  and  endoplasm,  as  is  common  among 
Protozoa.  The  cuticle  is  sometimes  regarded  as  the 
outer  portion  or  epicyte  of  the  ectoplasm.  A  single, 
vesicular,  spherical,  or  elliptical,  large  nucleus,  with 
its  chromatin  concentrated  to  form  a  spherical 
karyosome,  is  present.  The  body  of  some  gre- 
garines may  be  divided  by  ingrowing  ectoplasmic 
partitions  or  septa,  and  are  then  said  to  be  ^* septate"  or 
"polycystid"  (fig.  61).    Other  gregarines  remain  simple  and  non-septate. 


Fig.  59. — Monocystia 
agilis  from  seminal 
vesicles  of  Ltimbricus 

X  250.    (After Stein.) 


.[(;dl 


abed  e 

Fig.  60. — Gregaritia  longa  fiom  larva  of  crane-fly  {Tipula).     a,  in  epithelial  cell  of  host 
h,  f,  gradually  leaving  host-cell  ;  d,  adhering  to  host-cell  ;  e,  fully  developed  free  trophozoite. 


and  are  termed  'Mnonocystid  "  (fig.  59).  The  monocystid  gregarines 
occur  especially  in  the  body  cavity  of  Chaetopoda  and  Insecta,  more 
rarely    in    Echinodermata,   in  the    parenchyma  of    Platyhelminthes, 


GREGARINIDA 


-.  i^ 


also  in  the  gut  of  Tunicata  and  Insecta  (fig.  60)  and  in  the  seminal 

vesicles  of  Annelida.     In  the  polycystid  gregarines  a  single  septum  only 

is  present  as  a  rule,  and  thus  the  body  presents  two  portions :  (i)  an 

anterior  portion  termed  the  protomerite ;   (2)  a 

posterior,  larger  portion,  knOwn  as  the  deuto- 

merite,  which  generally  contains  the  nucleus. 

The  protomerite   is  often   modified  anteriorly 

to  form  an   organ  of  attachment,  termed  the 

epimerite  (fig.  61),  which  is  developed  from  the 

pointed  rostrum  of  the  sporozoite  or  primary 

infecting  young  gregarine.     The  structure   of 

the  epimerite  may  be  complicated,  being  pro- 
vided  with    hooks,   spines,  knobs,  and    other 

appendages.      An  extension  of  the  polycystid 

condition   is   seen    in   Tceniocystis   mira    Leger 

(from  the  dipteran  larva,  Ceratopogon  solstitialis), 

whose    body  shows   a   number   of   partitions, 

giving  the  organism  a  superficial  resemblance 

to  a  tapeworm. 

The  ectoplasm  of  a  gregarine  exhibits  three 

layers  :    (i)  An  epicyte  (cuticle)   externally  of 

which    the    epimerite    is     composed ;     (2)    a 

sarcocyte   which   forms  the  septa   if   present ; 

(3)  the  deeper  myocyte  layer  containing  con- 
tractile elements  in  the  form  of  fibrils  or  threads 

termed  myonemes  (fig.  62). 

The  endoplasm  is  fluid  and  granular,  con- 
taining many  enclosures,   which    are  of   the  nature   of   reserve  food 

materials.  They  consist  of  fat  drop- 
lets or  of  paraglycogen,  and  give 
the  organisms  an  opaque  appear- 
ance. Lithocystis  contains  crystals 
of  calcium  oxalate  in  its  endoplasm. 
Many  gregarines  are  capable  of 
active  movements,  though  they  do  not 
possess  obvious  locomotor  organs: 
The  movement  is  of  a  smooth, 
gliding  character  and  two  sug- 
gestions have  been  put  forward  to 
explain  it.  According  to  Schewiakoff, 
a  gelatinous  substance  is  secreted  be- 
tween the  layers  of  the  ectoplasm. 
This  is  extruded  posteriorly  and  thus 

the  animal  is  pushed  forward.     On  the  other  hand,  Crawley  consider^ 

that  the  movements  are  produced  by  contractions  of  the  myonemes. 


Fig.  61. — Xyphorhynchus 
firmus  with  epimerite  in 
intestinal  epithelial  cell  of 
host.     (After  Leger.) 


(i    E 

Fig.  62. — Gregarina  munieri  (from 
the  beetle,  Chrysomela  hcenioptera).  Sec- 
tion through  surface  layers.  C«,  cuticle  ; 
E^  ectoplasm  proper  ;  (7,  gelatinous 
layer  ;  My,  myonemes  in  myocyte  layer. 
X   1500.     (After  Schewiakoff.) 


132 


THE   ANIMAL    PARASITES   OF   MAN 


sporocyst ;  b,  mature  spore  con- 
taining sporozoites  ;  <r,  diagram- 
matic cross-section  of  spore, 
showing  eight  sporozoites  round 
residual  protoplasm.  (After 
Butschli.) 


These  two  explanations  are  probably  correct  as  far  as  each  goes,  and 
are  to  be  regarded  as  supplementary  to  one  another. 

Occasionally,  temporary  associations  of  gregarines  are  formed 
by  a  number  of  individuals  adhering  to 
one  another  end  to  end.  Such  temporary 
associations  are  examples  of  syzygy.  Such 
syzygies  must  not  be  confused  with  true 
associations  which  form  an  essential  part  of 
the  life-cycle. 

The  life-cycle  of  a  relatively  simple  gre- 
garine,  such   as   Monocystis   agilis    (fig.   59), 
parasitic  in  earthworms,  may  now  be  con- 
Monocystis  agilis.    sidered.     The  gregarines,  being  members  of 
Spores  from  vesicuia  seminalis   (he  Sporozoa,  produce  spores  at  One  phase 

of  the  Earthworm,    a,  Sporoblast       r     xi         rr  t  r^      i  • 

with  single  nucleus,  enclosed  in    o\    the    Jite-cycle.       li.ach    gregarme    spore 

(fig.  61)  develops  within  itself  a  number  of 
minute,  sickle-shaped  or  vermicular  bodies, 
known  as  sporozoites  or  primary  infecting 
germs.  Eight  sporozoites  are  often  formed 
within  each  spore.  When  absorbed  by  a  new 
host,  the  spore  softens  and  the  sporozoites  issue  from  it.  They  are 
capable  of  active  movement  and  may  or  may  not  enter  a  cell,  such  as 
one  of  those  of  the  digestive  tract,  or,  as  in  Monocystis,  a  cell  lining  the 
vesicuia  seminalis  which  becomes  a  sperm-cell  aggregate  (sperm  morula). 
When  the  sporozoite  has  reached  the  place  of  its  choice  in  the  host 
it  ceases  active  movements  and  proceeds  to  feed  passively  on  the  fluid 
substances  around  it,  whether  they  be  those  of  tissues  or  body  fluids. 
This  passive,  growing  and  feeding  form  is  known  as  the  trophozoite. 
After  a  trophic  existence  of  longer  or  shorter  duration,  the  trophozoite 
ceases  to  feed  and  prepares  for  reproduction.  Two  trophozoites 
associate  together,  each  of  them  first  becoming  somewhat  rounded. 
The  two  trophozoites,  now  known  as  sporonts  or  gametocytes,  become 
invested  in  a  single  common  envelope  or  cyst  (fig.  64,  a).  The  nucleus 
of  each  gametocyte  then  divides  by  a  series  of  binary  fissions  (fig.  64,  6), 
and  the  daughter  nuclei  thus  produced  arrange  themselves  at  the 
periphery  of  the  parent  cells  (fig.  64,  c).  Cytoplasm  collects  around 
each  of  these  nuclei,  and  thus  a  number  of  gametes  are  formed  within 
each  gametocyte.  The  gametes  for  a  time  exhibit  active  movements, 
and  ultimately  ripe  gametes  of  different  parentage  fuse  in  pairs,  that 
is,  conjugation  occurs  (fig.  64,  d).  In  this  way  zygotes  are  produced, 
the  nucleus  of  each  zygote  being  formed  by  the  fusion  of  two  gamete 
nuclei. 

The  zygote  grows  slightly  and  becomes  oval  or  elongate,  and  at 
this'period  is  often  called  the  sporoblast.     It  then  secretes  an  external 


GREGARINIDA 


133 


Fig.  64.— Schematic  figures  of  conjugation  and  spore  formation  in  Gregarines.     For  details 
see  text.     (After  Calkins  and  Siedlecki,  modified.) 


Fig.  65. — Stylorhynchtis  oblongatus.  or,  cyst  containing  two  sporonts  or  gametccytes,  each 
full  of  gametes,  those  in  the  upper  one  being  male,  b^  ripe  male  and  female  gametes,  x  1,600. 
(After  L.  Leger.) 


134 


THE   ANIMAL   PARASITES   OF   MAN 


membrane,  the  sporocyst.  Nuclear  division  occurs  inside  the  sporo- 
cyst  by  a  series  of  three  binary  fissions  (fig.  64,  e),  so  that  each  sporo- 
cyst, now  usually  referred  to  as  a  spore,  contains  eight  nuclei.  The 
cytoplasm  collects  around  each  nucleus  and  eight  vermicular  sporo- 
zoites  are  produced  within  each  spore  (Bg.  64,  /),  thus  completing 
the  life-cycle. 

It  will  be  noticed  that  in  the  above  life-cycle  no  asexual  multipli- 
cation occurs.  These  organisms,  such  as  Monocystis,  are  known  as 
the  Eugregarines,  and  include  the  majority  of  the  gregarines.  The 
remainder,  which  have  introduced  schizogony  into  their  life-cyle,  are 
known  as  the  Schizogregarines. 

There  are  variations  in  the  morphology  and  life-cycle  of  gregarines 
besides  those  that  have  been  mentioned.     It  is  not  within  the  scope 


Fig.  66. — Spores  of  various  Gregarines.  a,  Xipho- 
rhynchus.  b,  Ancyrophora.  c,  Gonospora.  d,  Cerato- 
spora.     (After  Leger.) 


of  this  book  to  discuss  them  in  detail,  but  the  following  may  be 
noted : — 

Morphological  differentiation  of  gametes  may  occur  as  in  S/y/o- 
rhynchus  oblongatus  (fig.  65),  which  differentiation  is  probably  of  a 
sexual  nature. 

The  sporocyst  really  consists  of  two  layers,  an  epispore  and  an 
endospore.  Externally  the  spores  of  different  gregarines  show  great 
variety  in  shape  and  markings,  and  spines,  or  long  processes  may 
be  present  (fig.  66). 

The  resistant  spores  serve  for  the  transmission  of  the  gregarines 
from  host  to  host.  The  mode  of  infection  is  contaminative,  the 
spores  expelled  with  the  dejecta  of  one  host  being  absorbed  with  the 
food  of  a  new  host. 

The  Gregarinida  may  be  classified  as  follows  : — 

Sub-order  I. — Eugregarinea,  without  schizogony. 

Tribe  i. — Acephalina. — Without  an  epimerite  and  non-septate  ; 
often    ^'  coelomic "    (body-cavity)    parasites.      E.g. :    Monocystis,   with 


COCCIDIIDEA  135 

several  speciesparasiti  c  in  the  seminal  vesicles  of  earthworms.  M^ny 
other  genera  parasitic  in  Echinodermata,  Tunicata,  Arthropoda,  etc. 

Tribe  2.~Cephalma. — With  an  epimerite,  either  temporarily  or 
permanently,  in  the  trophic  phase.  Usually  septate  (except  Dolio- 
cystidce).  Many  families,  genera  and  species.  Common  in  the  digestive 
tracts  of  insects.  E.g.  :  Gregarina,  with  several  species,  Gregarlna 
ovata  in  the  earwig,  Gregarina  hlattarum  in  the  cockroach,  Stylorhyn- 
chus  in  beetles,  Pterocephalus  in  centipedes,  etc. 

Sub-order  II. — Schizogregarinea,  with  schizogony. 

Tribe  i. — Endoschiza.^ — With  schizogony  occurring  in  the  intra- 
cellular phase,  e.g.y  Selenidium  (from  Annelida  and  Gephyrea),  Mero- 
gregarlna  (from  an  Ascidian).  , 

Tribe  2. — Edoschiza. — In  which  the  schizont  is  free,  and  schizogony 
is  extracellular,  e.g.,  Ophryocystis  (from  Blaps,  a  beetle),  and  Schizo- 
cystis  (from  Ceratopogon  larva). 

Order.     Coccidiidea. 

Hake  (1839)  first  saw  the  organisms  now  termed  Coccidia  during  his  investigations 
on  the  so-called  coccidial  nodules  of  rabbits.  The  opinions  as  to  the  nature  of  these 
peculiar  formations  were  very  diverse.  The  discoverer  considered  them  to  be  a  sort 
of  pus  corpuscle  ;  Nasse  (1843)  took  them  for  epithelial  cells  of  the  biliary  passages, 
others  for  helminthes,  especially  the  ova  of  trematodes  (Dujardin,  Kiichenmeister, 
Gubler,  etc  ).  Remak  (1845)  was  the  first  to  draw  attention  to  their  relation  to  the 
Psorospermia  (Myxosporidia),  and  this  investigator  found  them  also  in  the  small 
intestine  and  vermiform  appendix  of  rabbits.  Lieberkiihn  (1854),  who  examined  not 
only  the  coccidia  of  rabbits,  but  found  similar  forms  in  the  kidneys  of  frogs,  likewise 
called  them  definitely  psorosperms.  To  differentiate  Miiller's  psorosperms,  which  are 
found  in  fishes,  from  those  of  rabbits,  etc.,  the  latter  were  called  egg-shaped  psoro- 
sperms (Eimer),  until  R.  Leuckart  (1879)  named  them  Coccidia  and  placed  them  in 
a  group  of  the  Sporozoa  analogous  to  that  of  the  Gregarinida,  Myxosporidia,  etc. 
Numerous  works  confirmed  the  oQcurrence  of  coccidia,  not  only  in  all  classes  of 
vertebrate  animals,  but  also  in  invertebrates  (Mollusca,  Myriapoda,  Annelida,  etc.). 
A  large  number  of  genera  and  species  have  in  the  course  of  time  been  described  which 
inhabit  the  epithelium  of  the  intestine  and  its  appendages  for  choice,  but  are  also 
found  in  other  organs  (kidneys,  spleen,  ovaries,  vas  deferens,  testicles).  Some  also 
live  in  the  connective  tissue  of  various  organs,  more  particularly  of  the  intestine. 

The  knowledge  of  the  development  of  the  coccidia  was  of  particular  importance 
in  determining  their  classification.  By  means  of  encysted  coccidia  from  the  liver  of 
rabbits,  Kauffmann  (1847)  first  confirmed  the  fact  that  the  cyst,  which  was  partly  or 
entirely  filled  with  granular  contents,  divided  into  three  or  four  pale  bodies  (fig.  71) 
after  a  long  sojourn  in  water.  Lieberkiihn  observed  the  same  process  in  the  host  in 
the  case  of  the  coccidia  of  the  kidney  of  the  frog.  Stieda  (1865)  studied  more  minutely 
the  changes  that  occur  within  the  encysted  coccidia  of  the  liver  of  rabbits  after  the 
death  of  the  host.  He  discovered  that  the  bodies  now  known  as  "spores"  were 
oval  formations  pointed  at  one  pole,  and  surrounded  by  a  delicate  membrane,  which 
exhibited  a  certain  thickness  at  the  pointed  extremity  and  enclosed  a  slightly  bent 
rodlet,  expanding  at  either  end  into  a  strongly  light-refracting  globule  ;  a  finely 
granular  globule  was  present  in  the  middle  of  the  spore.   Waldenburg  (1862)  saw  the 

'  See  Fantham  (1908),  Parasitology,  i,  p.  369. 


136  THE    ANIMAL   PARASITES   OF   MAN 

same  phenomenon  in  coccidia  from  the  epithelium  of  the  villi  and  Lieberkiihn's 
glands  of  the  intestine  of  the  rabbit ;  but  the  process  in  this  case  took  place  in  a 
much  shorter  time. 

According  to  the  discovery  of  Kloss  (1855),  the  spores  of  the  coccidia  of  the 
urinary  organ  of  the  garden  snail  were  formed  in  far  greater  numbers  :  the  round 
spores  also  harboured  several  (five  to  six)  rodlets,  which  after  the  bursting  of  the 
spore-envelope  became  free.  Elmer's  researches  (1870)  afforded  information 
regarding  a  Coccidium  from  the  intestine  of  the  mouse,  which  was  transformed  in 
toto  into  a  "spore,"  containing  small  sickle-shaped  bodies.  The  fact  was,  moreover, 
established  that  the  little  bodies  left  the  delicate  envelope  when  in  the  intestine, 
made  movements  of  flexion  and  extension,  and  were  finally  transformed  into 
amoeboid  organisms,  which  apparently  penetrated  the  epithelial  cells  ;  at  all  events, 
similar  bodies  of  various  sizes  were  seen  in  these  cells.  Taking  the  immense 
number  of  these  parasites  into  account  and  the  lack  of  any  other  cause,  Eimer 
attributed  the  sudden  death  of  his  mice  to  the  Gregarina  falciformis,  as  the  parasite 
was  then  called,  just  in  the  same  way  as  a  few  years  previously  Reincke  ascribed  the 
acute  and  fatal  intestinal  catarrh  of  rabbits  to  the  invasion  of  intestinal  coccidia. 

All  that  had  become  known  about  coccidia  up  to  1879  was  then  compiled  by 
Leuckart,  and  completed  by  his  own  observations  on  the  coccidia  of  the  liver  of  the 
rabbit.  Experimental  infections  had  already  been  conducted  by  Waldenburg  (1862) 
with  intestinal  coccidia  of  rabbits,  and  by  Rivolta  (1869-73)  with  the  coccidia  of  fowls, 
which  experiments  confirmed  the  importance  of  the  spores,  or  bodies  enclosed  in  them, 
in  the  transmission  of  the  parasites  to  other  animals.  Accordingly,  it  was  assumed  that 
after  the  entry  of  the  spores  into  the  intestine  the  sporozoiles  were  set  free,  actively 
penetrated  into  the  intestinal  cells,  where  they  grew  into  coccidia,  and  finally  became 
encysted.  The  further  development,  i.e.^  the  formation  of  spores,  took  place  outside 
the  host's  body  in  these  cases  ;  in  other  cases  (Kloss,  Eimer)  it  took  place  within 
the  host.  Although  much  regarding  the  cycle  of  development  was  still  hypothetical, 
the  ideas  coincided  with  the  observations,  and  were  therefore  universally  regarded  as 
established.     Further  research  confirmed  this  view  in  numerous  new  forms. 

L.  Pfeiffer's  statements  (1891)  on  the  part  that  certain  coccidia  or  their  sporo- 
zoites  played,  or  seemed  to  play,  as  causes  of  disease  gave  a  renewed  impetus  to  the 
investigation  of  the  coccidia.  The  ingestion  of  even  very  numerous  spores  did  not 
appear  to  account  for  the  mass  infection  so  frequently  observed,  even  after  Balbiani 
had  confirmed  the  fact  that  there  were  two,  and  not  one,  sporozoites  contained  in  every 
spore  of  the  coccidia  of  rabbits  (fig.  72).  The  hypothesis  was  therefore  advanced 
that  the  sporozoites  or  young  coccidia  were  able  to  divide  once  again  by  sporulaiing. 
The  question  was  finally  solved  quite  differently.  R.  Pfeiffer  (1892)  first  confirmed 
the  fact  that  in  addition  to  the  well-known  method  of  sporulation  in  the  coccidia  of 
the  rabbit  that  causes  the  infection  of  fresh  hosts  ("exogenous  sporulation"),  an 
enormous  increase  may  follow  in  the  already  infected  host  in  a  manner  that  Eimer 
first  observed  in  the  coccidia  of  the  intestine  of  the  mouse  ("  endogenous  sporula- 
tion ").  It  had  hitherto  been  believed  that  some  of  the  species  of  coccidia  increased 
like  the  rabbit  parasite,  then  known  as  Coccidhun  ovifor?ne,  and  others  like  Eimeria 
falciformisy  and  this  difference  had  been  made  the  foundation  of  a  classification. 
R.  Pfeiffer  was  successful  in  observing  the  occurrence  of  both  kinds  of  development 
in  the  same  species,  and  expressed  the  opinion  that  endogenous  sporulation  (fig.  'j'^^ 
which  takes  place  within  the  host,  was  the  cause  of  the  mass-infection  that  is  mostly 
accompanied  by  serious  consequences  (fig.  74).  L.  Pfeiffer  sought,  especially,  to 
demonstrate  the  correctness  of  this  view  as  regards  other  species  of  coccidia 
and  for  this  purpose  he  utilized  the  exjoeriences  already  published.  Coccidia  were 
known  to  exist  in  a  number  of  different  hosts,  and  to  propagate  in  some  according 


COCCIDIIDEA 


137 


to  the  Coccidiutn  type,  in  others  according  to  the  Eimeria  type.  It  therefore  was 
reasoned  that  in  this  case  it  was  not  a  question  of  two  species  belonging  to  different 
genera  living  side  by  side,  with  a  different  mode  of  development,  but  of  one  species, 
in  the  life  of  which  both  forms  of  development  occurred  alternately. 

This  interpretation  of  facts  was  combated  especially  by  A.  Schneider  (1892)  and  by 
Labbe,  but  has,  nevertheless,  proved  true,  apart  from  the  circumstance  that  Schuberg 
succeeded  in  discovering  the  hitherto  unknown  Coccidium  form  in  the  intestine  of 
the  mouse ;  and  that,  moreover,  Leger  confirmed  the  fact  that  there  are  no  Arthro- 
poda  in  which  Eimeria  are  not  found  together  with  coccidia.  The  question  was 
finally  settled  by  experiments  made  by  Leger  with  the  coccidia  of  Scolopendra 
£ingulaia,  by  Schaudinn  and  Siedlecki  with  those  of  Lithobius  forjicatus^  and  by 
Simond  with  the  coccidia  of  the  rabbit.  On  Simond's  suggestion  the  sickle-shaped 
germs  corresponding  to  the  sporozoites,  which  are  formed  by  endogenous  sporula- 
tion,  are  generally  termed  merozoites  ;  and  in  accordance  with  Schaudinn's 
suggestion,  those  individuals  which  form  merozoites  are  termed  schizonts,  and 
those  which  produce  spores  are  called  sporonts.  In  contradistinction  to  sporogony 
(exogenous  sporulation),  the  term  schizogony  (endogenous  sporulation)  is  used. 

The  more  minute  examination  of  these  processes  at  last  led  to  the  discovery  of 
sexual  dimorphism,  of  copulation  and  of  alternation  of  generations  in  the  coccidia. 
Schuberg  was  the  first  to  consider  the  possibility  of  copulation  in  coccidia  ;  in 
addition  to  the  formations  which  now  are  termed  merozoites,  he  observed  very 
diminutive  bodies  ("  microsporozoites  ")  in  the  coccidia  of  the  intestine  of  the  mouse, 
which  were  able  eventually  to  copulate.  Labbe  confirmed  this  observation  in  some 
of  the  species,  and  Simond  expressed  the  opinion  that  bodies  termed  "chromato- 
zoites  "  occurred  in  all  coccidia.  Copulation  itself  was  then  observed  by  Schaudinn 
and  Siedlecki  (1897).  The  copulating  bodies  were  termed  gametes.  As,  however,  they 
differed  considerably  one  from  the  other,  the  males  were  called  microgametes  {i.e.^ 
the  microsporozoites  of  Labbe  and  the  chromatozoites  of  Simond)  and  the  females 
macrogametes.  After  copulation  was  completed  sporogony  took  place,  and  in  the 
cycle  of  development  of  one  species  this  regularly  alternated  with  schizogony  (asexual 
multiplication),  Schaudinn  in  1900  described  in  detail  the  life-cycle  of  Eimeria 
{Coccidium)  schubergi. 

The  recognition  of  this  unsuspected  complicated  process  was  bound  to  effect 
reforms  in  the  classification  of  the  coccidia  ;  and  all  the  forms  that  had  been 
regarded  as  developmental  stages  {Eimeria^  etc.)  had  to  be  reconsidered. 

Occurrence. — The  Coccidiidea  in  their  mature  condition  usually 
live  within  the  epithelial  cells  of  various  organs,  and  by  choice  inhabit 
those  of  the  intestine  and  of  its  associated  organs.  They  also  occur 
frequently  in  the  excretory  organs  of  mammals,  birds,  amphibia, 
molluscs,  arthropods,  and  may  also  be  found  in  the  testes  and  vas 
•deferens,  but  the  statement  that  they  hve  in  hen's  eggs,  as  well  as 
in  the  oviducts  of  fowls,  has  not  been  confirmed.^  Some  species 
inhabit  the  nuclei  of  cells,  others  live  in  the  connective  tissue,  but 

'  Notwithstanding  the  progress  made  during  the  last  decades,  the  ova  of  helminthes 
and  more  particularly  of  trematodes,  have  been  mistaken  for  Coccidia.  Thus  Poschinger 
{Zool.  Anz.,  1819,  ix,  p.  471)  and  Gebhard  {Virchoiv's  Arch.,  1897,  No.  147,  p.  536)  mistook 
the  ova  of  Distoma  turgidum,  Brds.,  for  Coccidia.  Podwyssotzki  {Centralbl.  f.  allg.  Path.., 
1890,  i,  p.  135)  made  a  similar  error  with  the  ova  (and  vitelline  sacs)  of  a  species  of  Prostho- 
^onimus  {Distoma  ovatum  of  the  authors)  ;  von  Willach  {Arch.  f.  wiss.  u.  prakt,  Thierheilk., 
1892,  xviii,  p.  242)  mistook  the  ova  of  a  nematode  for  Coccidia. 


138  THE   ANIMAL   PARASITES   OF   MAN 

their  presence  in  the  latter  situation  is  probably  only  secondary, 
that  is,  they  have  only  reached  it  from  the  epithelium  of  the  affected 
organs. 

The  size  of  the  Cocci diidea,  corresponding  as  a  rule  to  the  capacity 
of  their  habitat,  is  usually  small,  but  there  are  said  to  be  species 
that  attain  a  diameter  of  i  mm.  Their  form^  is  globular,  oval,  or 
elliptical.  External  appendages  are  lacking,  at  least  during  the  trophic 
or  vegetative  period  of  their  life,  which  is  spent  in  epithelial  cells, 
within  which  they  increase  in  size.  Frequently  one  only  is  present 
in  each  cell,  but  more  can  occur.  The  body  substance  is  composed 
of  a  more  or  less  finely  granular  or  distinctly  alveolar  protoplasm 
which  exhibits  no  differentiation  into  ecto-  and  endoplasm.  All 
species  possess  a  nucleus  that  enlarges  with  their  growth;  some- 
times it  only  shows  through  the  cytoplasm  as  a  lighter  spot,  or  may 
even  be  quite  concealed.  It  is  vesicular,  and  besides  containing  very 
delicate  threads  of  chromatin  in  the  clear  nucleoplasm,  it  contains 
generally  only  one  large  karyosome. 

-'The  infected  epithelial  cells  degenerate  sooner  or  later  as  the 
parasite  feeds  on  them  (fig.  67,  ii-iv).  After  their  form  has  been 
changed  by  the  action  of  the  growing  parasite,  they  finally  perish. 
The  cell  membrane  then  alone  surrounds  the  coccidia,  which,  at 
least  in  the  species  sufficiently  known,  begin  to  propagate  by  an 
asexual  process  (schizogony),  the  parasites  themselves  becoming 
schizonts,  as  the  initial  stage  is  usually  called.  They  differ  from  later 
stages  (sporonts  or  gametocytes),  which  resemble  them  in  form,  by 
the  absence  of  granulations  in  the  cytoplasm,  as  well  as  by  the 
vesicular  nucleus.  The  form  is  not  always  the  same,  for  in  some 
cases,  at  least,  many  schizonts  assume  a  globular  form. 

Schizogony  (fig.  67,  v-vii)  commences  with  a  division  of  the 
nucleus,  which  takes  place  in  different  ways  in  the  various  species. 
This  finally  leads  to  the  formation  of  numerous  daughter  nuclei 
which  become  smaller  and  smaller,  and  which  collect  beneath 
the  surface  of  the  schizonts.  In  some  species  the  daughter  nuclei 
collect  only  in  one  half  of  the  schizont.  A  part  of  the  protoplasm 
of  the  periphery  now  divides  around  each  daughter  nucleus,  the 
remaining  part  (residual  body)  being  left  in  the  centre  or  on  one 
side.  The  segments  of  the  divided  cytoplasm,  each  containing 
a  nucleus,  assume  a  fusiform  shape  and  become  merozoites,  which 
very  soon  become  free  (fig.  67,  viii)  and  leave  the  residual  body. 
They  are  distinguishable  from  the  very  similar  sporozoites  (fig.  67,  i), 
as  the  merozoites  possess  a  karyosome. 

*  The  life-cycle  given  here  is  based  on  that  of  Eimeria  [Coccidium)  schubergi,  after 
Schaudinn  (1900).  See  **  Untersuchungen  liber  den  Generationswechsel  bei  Coccidien,"  ZooU 
[ahrb.,  Abt.  f.  Anat.,  xiii,  pp.  197-292,  4  plates. 


COCCIDIIDEA 


^39 


Fig.  67. — Life-cycle  of  Eimeria  {Coccidiutn)  schubergi,  Schaud.,  from  the  intestine  of 
Liihobius.  (After  Schaudinn.)  The  infection  is  caused  by  a  cyst  (xx),  containing  spores, 
which  reaches  the  intestine  of  a  Lithobius,  where  it  discharges  the  sporozoites  (i).  11,  A 
sporozoite  invading  an  intestinal  epithelial  cell ;  iii,  intestinal  epithelial  cell  with  young 
trophozoite ;  iv,  intestinal  epithelial  cell  with  a  globular  schizont ;  v,  nuclear  segmentation 
within  the  schizont ;  vi,  the  daughter  nuclei  arranging  themselves  superficially  ;  vil,  forma- 
tion of  the  merozoites  ;  viii,  merozoites  that  have  become  free,  and  which,  penetrating  into 
other  epithelial  cells  of  the  same  intestine,  repeat  the  schizogony  (ii-viii) ;  ix  and  x, 
merozoites  which,  likewise  invading  the  epithelial  cells  of  the  same  intestine,  become 
sexually  differentiated  ;  xia,  young  macrogametocyte  ;  xib,  older  macrogametocyte ;  xic, 
mature  macrogametocyte  (discharging  particles  of  chromatin) ;  xiia,  young  microgametocyte  ; 
xiib,  older  microgametocyte  ;  xiic,  increase  of  nuclei  in  the  microgametocyte  ;  xiid,  the 
globular  residual  body  around  which  numerous  microgametes  have  formed  ;  xiie,  an  isolated 
microgamete  ;  xili,  the  mature  macrogamete  surrounded  by  numerous  microgametes  and 
forming  a  cone  of  reception  or  fertilization  prominence  ;  XIV,  shows  the  nucleus  of  a 
microgamete  that  has  penetrated  and  fused  with  the  nucleus  of  the  macrogamete  (fertilization) 
—the  latter  forms  a  membrane  and  becomes  an  oocyst ;  xv,  xvi,  xvii,  nuclear  segmentation 
in  the  oocyst;  xviii,  oocyst  with  four  sporoblasts ;  xix,  the  sporoblasts  transformed  into 
spores,  each  containing  two  sporozoites  ;  xx,  the  cyst  introduced  into  the  intestine  and 
liberating  the  sporozoites  by  bursting. 


140  THE   ANIMAL   PARASITES   OF   MAN 

The  merozoites  move  in  a  manner  similar  to  that  of  the  sporozoites.  The  move- 
ments consist  either  of  slow  incurvations  with  subsequent  straightenings,  or  annular 
contractions  along  the  entire  extent  of  the  body.  In  addition,  there  are  gliding 
movements  similar  to  those  of  many  gregarines,  and  brought  about  in  a  like  manner 
by  the  secretion  at  the  posterior  extremity  of  a  gelatmous  substance  that  hardens 
rapidly. 

The  merozoites  do  not  gain  the  open  in  the  usual  way,  but  are 
destined  to  infect  still  further  the  same  host  by  actively  penetrating 
into  other  epithelial  cells  of  the  affected  organ.  Here  they  continue 
their  growth  and  may  again  and  again  undergo  schizogony.  In  the 
Infusoria  the  repeated  segmentations  finally  cease  and  are  renewed 
only  after  a  conjugation.  This  is  likewise  the  case  with  the  Coccidia, 
with  the  difference  that  in  the  latter  the  two  conjugating  individuals 
(gametes)  are  differently  constituted  one  from  the  other,  whereas  in 
the  Infusoria  they  are  almost  always  similar. 

When  the  schizogony  ceases,  the  merozoites,  that  had  penetrated 
the  epithelial  cells  and  become  trophozoites  there,  consist  of  two  kinds 
of  differently  constituted  individuals.  One  kind  possesses  a  clear 
cytoplasm  (fig.  67,  Xli),  the  other  an  opaque,  richly  granular  cytoplasm 
(fig.  ()y,  xi),  while  both  possess  a  vesicular  nucleus  with  a  karyosome. 
In  order  to  continue  their  development,  the  more  granular  individuals 
must  be  fertilized,  and  are  therefore  termed  either  female  gametes  or, 
on  account  of  their  size,  macrogametes.  The  male  individuals  (micro- 
gametes)  necessary  to  conjugation,  are  formed  in  greater  numbers 
from  the  less  dense  microgametocytes  or  male  mother-cells  (fig.  67, 
xiid).  They  are  slender  bodies  consisting  chiefly  of  nuclear  substance, 
and  in  most  species  bear  two  flagella  of  unequal  length  directed  back- 
wards, the  place  of  insertion  of  which  varies  according  to  the  species 
(fig.  67,  xiie). 

While  the  development  of  the  microgametes  is  rapidly  advancing 
a  change  occurs  in  the  nucleus  of  the  female  parent  forms  or  macro- 
gametocytes.  Parts  of  the  karyosome  are  extruded  (fig.  67,  xic),  and 
the  nucleus  loses  at  the  same  time  its  vesicular  form.  One  macro- 
gamete  results,  after  nuclear  maturation,  from  one  macrogametocyte. 
By  this  time  the  macrogametes  are  capable  of  conjugation,  and 
the  process  takes  place  within  the  host,  generally,  however,  outside 
the  affected  and  degenerated  host  cells.  The  microgametes  that 
have  now  become  free  from  the  very  large  residual  body,  crowd 
around  the  mature  macrogametes,  which  often  send  out  a  small 
prominence  ("cone  of  reception"  or  fertilization  protuberance)  for 
their  reception  (fig.  67,  xiii).  As  soon  as  a  microgamete  comes  in 
contact  with  this  and  penetrates  into  the  cytoplasm  of  the  macro- 
gamete,  the  latter  surrounds  itself  with  a  membrane  which  prevents 
the  intrusion   of  other   microgametes.      The    nucleus  of  the  micro- 


COCCIDIIDEA  141 

gamete  that  has  gained  entry  unites  with  the  nucleus  of  the  macro- 
gamete,  which  latter  is  afterwards  capable  of  forming  the  well-known 
spores.  The  parasite  is  now  called  an  encysted  zygote  or  oocyst.  The 
oocysts  of  some  other  members  of  the  Coccidiidea,  e.g.,  Ehuerla  avium, 
can  form  their  walls  prior  to  fertilization.  In  such  cases,  a  weak  spot 
is  left  at  one  place  in  the  cyst  wall,  forming  a  micropyle,  that  permits  of 
the  entry  of  the  male,  immediately  after  which  the  micropyle  is  closed. 

The  reduced  nucleus  of  the  macrogamete  elongates  on  the  entry  of  the  micro- 
gamete,  and  becomes  a  fertilization  spindle  to  which  the  male  pronucleus  (from  the 
microgamete)  becomes  attached  (fig.  67,  xiv  and  xv).  Thereupon  the  spindle 
divides  into  two  daughter  nuclei  (fig.  67,  xvi)  which  assume  a  round  shape.  The 
protoplasm  at  this  stage  may  at  once  divide,  or  another  segmentation  of  the  daughter 
nuclei  may  first  occur.  In  the  former  case  the  two  halves  divide  again,  so  that 
finally  four  nucleated  segments,  the  sporoblasts,  are  formed,  whereas  in  the  latter 
case  the  four  sporoblasts  appear  simultaneously  (fig.  67,  xvn).  In  both  cases  a 
residual  body  of  varying  size  is  separated  from  the  protoplasm  of  the  oocyst.  As 
a  rule  the  oocysts  have  already  been  discharged  from  the  body  of  the  host,  and  in 
the  manner  described  above,  form  the  sporoblasts  after  a  longer  or  shorter  period 
of  incubation. 

The  sporoblasts  are  originally  naked,  but  each  soon  secretes  a 
homogeneous  membrane,  the  sporocyst,  in  which  it  becomes  enveloped 
(fig.  67,  xviii).  After  the  segmentation  of  the  nucleus  the  contents 
divide  into  two  sickle-shaped  sporozoites,  in  addition  to  which  there 
is  generally  also  a  residual  body  (fig.  67,  xix). 

This  terminates  the  development.  The  spores  are  intended  for 
the  infection  of  other  hosts.  If  they  reach  the  intestine  of  suitable 
hosts,  either  free  or  enclosed  in  the  oocyst  wall,  the  action  of  the 
intestinal  juices  causes  them  to  open  and  permits  the  escape  of  the 
sporozoites  (fig.  67,  xx).  The  latter  move  exactly  like  the  merozoites 
and  soon  make  their  way  into  epithelial  cells  (fig.  67,  i),  where  they 
become  schizonts,  and  thus  repeat  the  life  cycle. 

Although  our  knowledge  of  the  development  of  the  coccidia  is  but  of  recent  date, 
yet  it  already  extends  to  a  large  number  of  species,  which  exhibit  various  deviations 
from  the  cycle  of  development  described  above.  For  instance,  in  addition  to 
differences  in  the  gametocytes,  the  schizonts  of  Adelea  and  Cyclospora  also  show 
differentiation  and  give  rise  to  macromerozoites  and  micromerozoites,  whilst  in 
Adelea  and  Klossia  a.  precocious  association  of  the  gametocytes  precedes  the  true 
copulation  of  the  ripe  gametes. 

The  classification  of  the  Coccidiidea  is  based  chiefly  on  the 
number  of  sporozoites  found  in  each  spore,  and  the  number  of 
sporocysts  (spores)  found  in  one  oocyst.  Leger'  recognises  two 
great  legions,  the  Eimeridea  and  the  Adeleidea,  the  former  comprising 
the  greater  number  of  genera,  including  the  genus  of  most  economic 
importance,  Eiiueria.     It  must  be  noted  that,  though  a  member  of 

'  Arch.f.  Frotistenktmde  (191 1),  xxii,  p.  71. 


142  THE   ANIMAL   PARASITES    OF   MAN 

this  genus  may  be  frequently  referred  to  as  Coccidinniy  strictly  it 
should  be  termed  Eimeria,  that  name  having  priority.  The  name 
of  the  disease  resulting  from  the  action  of  such  parasites  is,  however, 
established  and  remains  as  coccidiosis. 

Certain  of  the  more  important  of  the  Coccidiidea  may  now  be 
considered. 

Genus.    Eimeria,  Aime  Schneider,  1875. 

Syn. :  Psorospermiuni,  Rivolta,  1878  ;  Cytospermiiim,  Rivolta,  1878  ;  Coccidiiim^ 
R.  Leuckart,  1879;  Pfeifferia,  Labbe,  1894:   Pfeifferella,  Labbe,   1899. 

The  Eimeria  belong  to  Leger's  old  family,  the  Tetrasporocystidae, 
which  comprises  forms  producing  oocysts  with  four  sporocysts,  each 
containing  two  sporozoites.  The  cysts  are  spherical  or  oval,  as  are 
also  usually  the  schizonts.  The  members  of  the  genus  are  confined 
chiefly  to  vertebrate  hosts,  the  more  important  economically  occurring 
in  mammals  and  birds.  From  the  mammalian  hosts  very  rarely  the 
parasites  may  reach  man.  Eimeria  (Coccidium)  avium  of  wild  birds 
and  poultry,  and  Eimeria  stieda^  parasitic  in  rabbits,  may  be  con- 
sidered. There  is  a  general  similarity  in  their  life-cycles  and  each  is 
of  great  practical  importance. 


Eimeria  avium,  Silvestrini  and  Rivolta. 

Eimeria  avitmi  is  responsible  for  fatal  epizootics  among  game  birds  such  as 
grouse,  pheasants  and  partridges,  and  domestic  poultry  such  as  fowls,  ducks,  pigeons 
and  turkeys,  and  can  pass  from  any  one  of  these  hosts  to  any  of  the  others  with  the 
same  effect.  The  organism  is  parasitic  in  the  alimentary  tract  of  the  host,  affecting 
more  ejJpWBcially  the  small  intestine  (duodenum)  and  the  casca,  but  in  some  cases 
penetrating  to  the  liver  and  multiplying  there  (as  in  turkeys),  producing  necrotic 
cheesy  patches,  that  ultimately  become  full  of  oocysts.  The  gut  is  rendered  very 
frail  by  the  action  of  the  parasites,  its  mucous  membrane  is  greatly  injured,  and 
is  often  reduced  to  an  almost  structureless  pulp,  riddled  with  parasites  (fig.  68). 
Infection  is  conveyed  from  host  to  host  by  the  ingestion  of  food  or  drink  contamin- 
ated with  the  oocysts  voided  in  the  fieces  of  infected  birds.  Oval  oocysts  from  24  /^  to 
35  ;u  long  and  from  14 /x  to  20  ^u  broad  are  the  means  of  infection.  The  oocysts 
develop  internally  four  sporocysts  or  spores,  from  each  of  which  two  sporozoites  aie 
produced.  The  life-history^  presents  two  phases:  (i)  The  asexual  multiplicative 
phase,  schizogony,  for  the  increase  in  numbers  of  the  parasites  within  the  same  host  ; 
(2)  the  reproductive  phase,  following  the  formation  of  gametes  (gametogony),  leading 
to  the  production  of  resistant  oocysts,  destined  for  the  transference  of  the  parasite 
to  new  hosts  (sporogony). 

The  oocysts  usually  reach  the  duodenum  unharmed,  with  food  or  drink.     Under 

'  Fantham,  H.  B.  (1910),  "The  Morphology  and  Life  History  of  Eimeria  (Coccidiwn) 
avium,  a  Sporozoon  causing  a  fatal  disease  among  young  Grouse,"  Proc.  Zool.  Soc.  Lend., 
1910,  pp.  672-691,  4  plates.  Also  Fantham,  H.  B.  (1911),  "Coccidiosis  in  British  Game 
Birds  and  Poultry,"  Joiirn.  Econ.  Biol.,  vi,  pp.   75-96. 


EIMERIA   AVIUM 


143 


the  influence  of  the  powerful  digestive  juices  (especially  the  pancreatic)  now  en- 
countered, the  oocysts  soften,  as  do  the  sporocysts,  and  ultimately  two  sporozoites 
emerge  from  each  sporocyst.  The  sporozoites  are  from  y  fi  to  10  /j.  long,  and  each  is 
vermicular  with  a  uniform  nucleus  (fig.  69,  a).  After  a  short  period  of  active  move- 
ment in  the  gut,  each  sporozoite  penetrates  an  epithelial  cell  (figs.  68  spz,  69,  b),  and 
once  within,  gradually  becomes  rounded  (fig.  69,  B,  c).  It  grows  rapidly,  feeding  on 
the  contents  of  the  host  cell  and  living  as  a  trophozoite  (fig.  69,  d).  When  the  parasite 
is  from  10  a*  to  12  /t  in  diameter,  usually  multiplication  by  schizogony  (fig.  69,  e-h) 
begins.  The  nucleus  of  the  parent  cell,  now  called  a  schizont,  divides  into  a  number 
of  portions  that  become  arranged  at  the  periphery  (fig.  69,  e).  Cytoplasm  collects 
around  each  nucleus  (fig.  69,  E,  f)  and  gradually  a  group  of  daughter  individuals 
(merozoites)  is  produced  (fig.  69,  G),  the  nucleus  of  each  merozoite  showing  a 
karyosome. 

The  merozoites  oi Ei77ieria  aviuin  are  arranged  "en  barillet,"  like  the  segments  of 


r--^v.r  V  ^  ®  ''^  V 


'v«»3 


Fig.  68. — Small  piece  of  epithelial  lining  of  gut  of  heavily  infected  Grouse  chick,  show- 
ing various  stages  in  life  history  of  the  parasite  Einieria  avium  ;  par^  parasite  (trophozoite) ; 
mz^  merozoite  ;  sch^  schizont ;  jt/s,  sporozoite  ;  ooc^  oocyst ;  cT,  male  gametocyte  ;  ?  ,  female 
gametocyte.      X  750.     (After  Fantham.) 


an  orange  (figs.  68  mz^  69,  G),  therein  difTering  from  those  of  E.  schubergi,  which  are 
arranged  "  en  rosace."  They  separate  from  one  another  (fig.  69,  H),  penetrate  other 
epithelial  cells,  where  they  may,  in  turn,  become  schizonts.  Eight  to  fourteen  mero- 
zoites are  usually  formed  by  each  schizont,  twenty  have  been  found,  while  in  cases 
of  intense  infection  when  space  has  become  limited,  the  number  may  be  only  four. 

After  a  number  of  generations  of  merozoites  have  been  formed,  a  limit  is  reached 
both  to  the  multiplicative  capacity  of  the  parasite  and  to  the  power  of  the  bird 
to  provide  the  invader  with  food.  Consequently,  resistant  forms  of  the  parasite 
are  necessary,  and  the  trophozoites  begin  to  show  sexual  dififerentiation  instead 
of  forming  schizonts,  that  is,  gametogony  commences. 

Certain  trophozoites  store  food  and  become  large  and  granular.  These  are 
macrogametocytes  (fig.  69,  i,  $).  The  microgametocytes  (fig.  69,  i,  ^)  are 
smaller   and  far   less   granular.     The   macrogametocyte    continues    to   grow,    and 


144 


THE   ANIMAL   PARASITES   OF   MAN 


becomes  loaded  with  chromatoid  and  plastinoid  granules  (fig.  69,  j,  ? ),  while 
the  microgametocyte  has  its  nucleus  divide  to  form  a  number  of  bent,  rod-like 
portions  (fig.  69,  J,  S)-  The  macrogametocyte  gives  rise  to  a  single  macrogamete, 
which  forms  a  cyst  wall  for  itself,  leaving  a  thin  spot  (micropyle)  for  the  entry 
of  the  male  (fig  69,  K,  ? ).  The  microgametocyte  gives  rise  to  numerous  small, 
biflagellate  microgametes  (fig.  69,  K,  ^)  around  a  large,  central  residual  mass, 
from  which  they  ultimately  break  free,  and  swim  away.  When  a  macrogamete 
is  reached,  the  microgamete  enters  through  the  micropyle  (fig.  69,  l) — which 
then    closes,    thus   excluding   the    other   males — and   applies    itself  to   the  female 


Fig.  69. — Eimeria  avium.     Diagram  of  life-cycle.     For  explanation  see  text. 
(After  Fantham.) 


nucleus  (fig.  69,  m).  Nuclear  fusion  occurs,  the  oocyst  (encysted  zygote)  being  thus 
produced.  Sporogony  then  ensues.  The  oocyst  (fig.  69,  N)  at  first  has  its  contents 
completely  filling  it.  They  then  concentrate  into  a  central  spherical  mass  (fig.  69,  o) 
which  gradually  becomes  tetranucleate  (fig.  69,  p).  Cytoplasm  collects  around  each 
nucleus,  and  four  sporoblasts  are  thus  formed  (fig.  69,  q).  Each  sporoblast  becomes 
oval  (fig.  69,  r)  and  produces  a  sporocyst.  Ultimately  two  sporozoites  are  formed  in 
each  sporocyst  or  spore,  at  first  lying  tete-beche  (fig.  69,  s),  but  finally  twisting 
to  assume  the  position  most  convenient  for  emergence  (fig.  69,  t)  when  they  reach  a 
new  host.     The  period  of  the  life-cycle  of  Eimeria  avium  (as  well  as  the  details 


EIMERIA   STIED^ 


145 


of  the  life-cycle)  was  determined  by  Fantham  to  be  from  eight  to  ten  days,  of  which 
period  schizogony  occupies  four  to  five  days. 

The  method  of  infection  ^  is  contaminative,  by  way  of  food  or  drink.  Young 
birds  are  especially  susceptible  to  infection.  Certain  birds,  particularly  older  ones, 
may  act  as  reservoirs  of  oocysts,  being  continuously  infected  themselves,  without 
showing  any  marked  ill  effects  from  the  parasite,  but  being  highly  infectious 
to  other  birds.  Much  moisture  retards  the  development  of  sporocysis  consider- 
ably. The  duration  of  vitality  of  the  infective  oocysts  has  been  determined 
experimentally  to  extend  well  over  two  years,  and  in  certain  cases  longer. 
Eiineria  avium  is  the  causal  agent  of  "white  diarrhoea"  or  "white  scour"  in 
fowls,  and  of  "  blackhead  "  in  turkeys. 

Euneria  avium  of  birds  and  E.  stiedce  of  rabbits  closely  resemble 
one  another,  but  are  not  the  same  parasite,  for  E,  avium  is  not  infective 
to  rabbits,  nor  E.  stiedce  to  poultry. 

Eimeria  stiedae,   Lindemann,  1865. 

Syn. :  Monocystis  stiedce^  Lindemann,  1865;  Psorospermium  cunicult,  Rivolta, 
1878;  Cytospermium  homi?tis^  Rivolta,  1878;  Coccidium  oviforrne^  Leuckart,  1879; 
Coccidium  perforans^  Leuckart,   1879;  Coccidium  cuniculi. 

Eimeria  stiedcv.  is  parasitic  in  the  gut  epithelium  (fig.  70),  liver,  and 
epithelium  of  the  bile  ducts  of  rabbits,  and  is  usually  considered  to 


Fig.  70. — Eimeria  stiedce.     Section  through  an  infected  villus  of  rabbit's  intestine.      X  260. 

be  the  parasite  very  occasionally  found  in  man.  The  life-cycle 
resembles  that  of  Eimeria  avium  in  its  general  outlines  (see  fig.  69) 
and  therefore  will  not  be  detailed  in  full  here.  The  oocysts  (fig.  71) 
are  large,  elongate-oval,  greenish  in  fresh  preparations  and  vary  in 
size  from  24  yu,  to  49  //<  long  and  12*8  //,  to  28  //,  broad,  the  gut  forms 
being  usually  smaller  than  those  occurring  in  the  liver,  owing  to  the 


'  Fantham,  H.   B.   (1910),   '*  Experimental  Studies  on  Avian  Coccidiosis,   especially   in 
relation    to   young   Grouse,   Fowls   and    Pigeons,"    Proc.  Zool,   Soc.    Land.,    1910,  pp.   722- 
731,   I   plate. 
10 


146 


THE    ANIMAL   PARASITES   OF   MAN 


more  confined  space  in  which  they  are  formed.  Formerly,  the  para- 
sites in  the  Hver  were  described  under  the  name  of  Coccidium  oviforme, 
while  those  from  the  intestine  were  termed  Coccidiiun  perforans.  This 
distinction  has  now  broken  down. 

The  oocysts^  are  thick-walled,  somewhat  flattened  at  one  pole, 
where  a  large  micropyle  is  present.  Four  egg-shaped  spores  (sporo- 
cysts)  are  formed  within,  each  about  12  //,  to  15  yLt  long  and  7  yit  broad 


Fig.  71. — Eimeria  stiedce,  from 
the  liver  of  the  rabbit,  oocysts 
in  various  stages  of  development. 
(After  Leuckart.) 


Fig.  72.— a,  b,  spores  of 
Eimeria  stiedce  (Riv.),  with 
two  sporozoites  and  residual 
bodies ;  c  represents  a  free 
sporozoite.    (After  Balbiani.) 


(fig.  72).  The  oocysts  are  voided  with  the  faeces.  Sporogony  takes, 
in  nature,  about  three  days  in  the  excrement.  Faecal  contamination 
of  the  food  of  rabbits  results,  and  coccidian  oocysts  are  swallowed. 
Under  the  influence  of  the  pancreatic  juice  of  a  new  host,  the 
sporozoites  (fig.  72,  a — c)  are  liberated  from  the  spores  and  proceed 
to  attack  the  epithelium  and  multiply  within   it,   as  in    the  case  of 

Eimeria  aviiiin.     From  the  gut, 
infection    spreads    to    the    liver, 
where  multiplication  of  the  para- 
site  goes    on    actively,  resulting 
in  the  formation  of  the  whitish 
coccidial  nodules,  which  may  be 
very  conspicuous  (fig.  74).     Pro- 
liferation of  the  connective  tissue 
may  occur  around  the  coccidial 
nodules,  which  then  contain  large 
numbers    of   oocysts   in   various 
stages  of  development.     It  is  said  that  the  oocysts  in  the  older  nodules 
do  not  seem  to  be  capable  of  further  development.    Schizogony  (fig.  73) 
and  gametogony  in  all  stages  can  be  found  in  both  liver  and  gut. 


Fig.  73. — So-called  swarm  cysts  (endogenous 
sporulation  or  schizogony)  of  the  Coccidium  of 
the  rabbit.  The  daughter  forms  are  called 
merozoites.     (After  R.  Pfeiffer.) 


*  For  an  account  of  the  life-cycle  of  Eimeria  stiedce  consult  Wasielewski,  Th.  von  (1904), 
*'Studien  und  Photogramme  zur  Kenntnis  der  pathogenen  Protozoen,"  Heft.  I  (Coccidia), 
118  pp.,  7  plates,  Leipzig:  J.  A.  Earth.    Also,  Metzner,  R.  [\f^oi)^  Arch.  f.  Frotistenk.,\\,T^.  13. 


EIMERIA   STIED/E 


147 


Young  rabbits  often  die  of  intestinal  coccidiosis  before  infection 
of  the  liver  occurs.  The  repeated  schizogony  of  Euiieria  stiedcv  in  the 
gut  is  sufficient  to  cause  death. 


li: 

^1 

) 

tvSk^ 

i'^S 

m- 

Fig.  74. — Eimeria  stiedce.     Section  through  coccidian  nodule  in  infected  rabbit's  liver,     x  5S« 


The  disease  of  cattle  popularly  known  as  "  red  dysentery  "  is  also  ascribed  to  the 
action  of  Eimeria  stiedce.  The  faeces  of  infected  cattle  show  blood  clots  of  various 
sizes  and  in  severe  cases  watery  diarrhoea  is  present.  Acute  cases  end  fatally  in 
about  two  days.  Numerous  oocysts,  considered  to  be  those  oi  Eiineria  stiedce.,  occur 
in  the  faeces,  and  there  is  a  heavy  infection  of  the  gut,  especially  the  large  intestine 
and  rectum,  all  stages  of  the  parasite  being  found  in  the  epithelium.  It  is 
suspected  that  cattle  contract  the  disease  by  feeding  on  fresh  grass  contaminated 
with  oocysts.     The  disease  is  recorded  from  Switzerland  and  from  East  Africa. 


148  THE   ANIMAL   PARASITES    OF   MAN 

As  before  mentioned,  Einieria  stiedce  is  considered  to  be  the 
organism  found  in  a  few  cases  in  man,  possibly  acquired  by  eating 
the  insufficiently  cooked  livers  of  diseased  rabbits.  These  cases  may 
now  be  described. 

{a)  Human  Hepatic  Goccidiosis. 

(i)  Gubler's  Case.  A  stone-breaker,  aged  45,  was  admitted  to  a  Paris  hospital 
sufifering  from  digestive  disturbances  and  severe  anaemia.  On  examination  the  liver 
was  found  to  be  enlarged  and  presented  a  prominent  swelling,  which  was  regarded  as 
being  due  to  Echinococcus.  At  the  autopsy  of  the  man,  who  succumbed  to  intercurrent 
peritonitis,  twenty  cysts  were  found  averaging  2  to  3  cm.  in  diameter,  and  one 
measuring  12  to  15  cm.  The  caseous  contents  consisted  of  detritus,  pus  corpuscles, 
and  oval-shelled  formations,  which  were  considered  to  be  Distoma  eggs,  but  which, 
in  accordance  with   Leuckart's  conjecture,  proved  to  be  Coccidia.^ 

(2)  Dressler's  Case  (Prague).  Relates  to  three  cysts,  varying  from  the  size  of 
a  hemp-seed  to  that  of  a  pea,  and  containing  Coccidia,  found  in  a  man's  liver.  =* 

(3)  Sattler's  Case  (Vienna).  Coccidia  were  in  this  case  observed  in  the  dilated 
biliary  duct  of  a  human  liver.'' 

(4)  Perls'  Case  (Giessen).  Perls  discovered  Coccidia  in  an  old  preparation  of 
Sommering's  agglomerations.'' 

(5)  Silcock's  Case  (London).^  The  patient,  aged  50,  who  had  fallen  ill  with 
serious  symptoms,  exhibited  fever,  enlarged  liver  and  spleen,  and  had  a  dry,  coated 
tongue.  At  the  autopsy  numerous  caseous  centres,  mostly  immediately  beneath  the 
surface,  were  found,  while  the  contiguous  parts  of  the  liver  were  inflamed.  Micro- 
scopical examination  demonstrated  numerous  Coccidia  in  the  hepatic  cells  as  well 
as  in  the  epithelium  of  the  biliary  ducts.  A  deposit  of  Coccidia  was  likewise  found 
in  the  spleen,  which  the  parasites  had  probably  reached  by  means  of  the  blood- 
stream.*^ 

{h)  Human  Intestinal  Goccidiosis. 

In  two  cadavers  at  the  Pathological  Institute  in  Berlin,  Eimer^  found  the 
epithelium  of  the  intestine  permeated  by  Coccidia.  Railliet  and  Lucet's  case  may 
be  traced  back  to  intestinal  Coccidia,  which  were  found  in  the  faeces  of  a  woman 
and  her  child,  who  had  both  suflfered  for  some  time  from  chronic  diarrhoea.^  In 
other  cases  (^Grassi,  Rivolta),  where  only  the  existence  of  Coccidia  in  the  faeces  was 
known,  it  is  doubtful  whether  the  parasites  originated  in  the  intestine  or  in  the  liver. 

'  Gubler,  A.,  "  Tumeurs  du  foie  determ.  par  des  oeufs  d'helm  .  .  .  ."  Me>?i.  Soc. 
Biol.,  Paris,  1858,  v,  2  ;  and  Gaz.  med.  de  Paris,  1858,  p.  657  ;  Leuckart,  R.,  Die  menschl. 
Paras.,  1863,  ist  edition,  i,  pp.  49,  740. 

^  Leuckart,  R.,  Die  menschl.  Paras.,  1863,  1st  edition,  i,  p.  740.- 

^  Leuckart,  R.,  Die  Pai-as.  d.  mensch.,  1879,  2nd  edition,  p.  281. 

*  Leuckart,  R.,  ibid.,  p.  282. 

^  Silcock,  "A  Case  of  Parasit.  by  Psorospermia,"  Trans.  Path.  Soc,  London,  1890,  xli, 
p.  320. 

*  Pianese  has  confirmed  the  fact  that  Coccidia  actually  occur  in  the  blood  of  the  hepatic 
veins  of  infecteil  rabbits. 

'  Die  ei-  ti.  kiigelf.  Psorosp.  d.  Wirbelt.,  1870,  p.  16. 

*  Railliet  and  Lucet,  "  Obs.  s.  quelq.  Cocc.  intest.,"  C.  R.  Soc.  Biol.,  Pari?,  1890,  p.  660  ; 
Railliet,  Trait.  Zool.  med.  et  agric,  2e  ed.,  1895,  p.  140. 


ISOSPORA    BIGEMINA  149 

(c)  Doubtful  Cases. 

To  these  belong  Virchow's  case  '  where,  in  the  liver  of  an  elderly  woman,  a  thick 
walled  tumour  measuring  9  to  1 1  mm.  was  found.  Among  the  contents  of  this 
tumour  there  were  oval  formations  56  fi  long,  surrounded  by  two  membranes  and 
enclosing  a  number  of  round  substances.  Virchow  considered  these  foreign  bodies 
to  be  eggs  of  pentastomes  in  various  stages  of  development,  others  consider  them 
to  be  Coccidia. 

The  Coccidia  which  Podwyssotzki  claims  to  have  seen  in  the  liver  of  a  man,  not 
only  in  the  liver  cells,  but  also  in  the  nuclei,  are  also  problematic.^  The  parasite 
was  called  Caryophagus  hominis. 

Again,  other  explanations  can  be  given  to  an  observation  by  Thomas,  on  the 
occurrence  of  Coccidiuin  oviforme  in  a  cerebral  tumour  of  a  woman  aged  40.  The 
growth  was  as  large  as  a  pea  and  surrounded  by  a  bony  substance.'^ 

Genus.     Isospora,  Aime  Schneider,  1881. 

Syn. :  Dipiospora^  Labbe,  1893. 

Belonging  to  the  section  Disporea^  that  is,  forming  only  two  spores, 
each  with  four  sporozoites. 

Isospora  bigemina,  Stiles,  1891. 

Syn.:    '''' Cytospermiuiti  villoru7ii   i?itestinalium   ca?tis  et  felis^''  Rivolta,   1874; 
"  Coccidium  Rivolta^''  Grass),  1882  ;    Coccidium  bigeininum,  Stiles,  1891. 

This  parasite  lives  in  the  intestinal  villi  of  dogs,  cats,  and  the 
polecat  (Mustela  putorins,  L.).  According  to  Stiles,''  the  oocyst 
divides  into  two  equal  ellipsoidal  portions  or  sporoblasts  which 
become  spores  and  then  each  forms  four  sporozoites.  The  oocysts 
of  this  species  vary  from  22  /x,  to  40  yu,  in  length  and  from  19  fju  to 
28//.  in  breadth.  Each  spore  is  10//,  to  18 /^  long  and  contains 
four  sporozoites.  The  parasites  live  and  multiply,  not  only  in  the 
gut  epithelium,  but  also  in  the  connective  tissue  of  the  intestinal 
submucosa.     Wasielewski  has  seen  merozoites  in  the  gut  of  the  cat. 

Isospora  bigemina  (fig.  75)  appears  to  occur  also  in  man,  for 
Virchow  published  a  case  which  was  communicated  to  him  by 
Kjellberg,  and  attributed  the  illness  to  this  parasite.^  Possibly  also  it 
would  be  more  correct  to  ascribe  the  observation  of  Railliet  and 
Lucet,  which  is  mentioned  under  "  Human  Intestinal  Coccidiosis," 
p.  148,  to  this  species,  as  the  Coccidia  in  that  case  were  distinguished 

'  Arch,  f.  path.  An.^  xviii,  i860,  p.  523. 

2  Podwyssotzki,  "  Ueb.  d.  Bedeut.  d.  Coccid.  in  d.  Path.  Leber  des  Menschen,"  Centralbl. 
f.  Bakt.,  vi,  1889,  p.  41. 

3  Thomas,  J.,  "Case  of  Bone  Formation  in  the  Human  Brain,  due  to  the  Presence  of 
Coccidium  oviforme,''  [ournal  Boston  Soc.  Med.  Sc,  iii,  1899,  p.  167;  Centra/d/.  /.  Bakt. 
[ij  xxviii,  1900,  p.  882. 

*  *'  Notes  on  Paras.,"  No.  11,  /ourn.  of  Comp.  Med.  and  Vet.  Sci.,  1892,  xiii,  p.  517. 

*  Arch,  f  path.  An.,  i860,  xviii,  p.  527. 


ISO 


THE   ANIMAL   PARASITES   OF   MAN 


by  their  diminutive  size  (length  15  //,,  breadth  10  /x).     The  case  com- 
municated by  Grunow  may  also  possibly  refer  to  hospora  bigemina} 


Fig.  75. — hospora  bigemina,  Stiles  (from  the  intestine  of  a  dog),  a,  Piece  of  an  intestinal 
villus  beset  with  Isospora,  slightly  enlarged  ;  b,  Isospora  bigemina  {15  fi  in  diameter),  shortly 
before  division  ;  c,  divided  ;  d,  each  portion  encysted  forming  two  spores  ;  ^,  four  sporozoites 
in  each  part,  on  the  left  seen  in  optical  section, together  with  a  residual  body— highly  magnified. 
(After  Stiles.) 

Roundish  or  oval  structures  of  6  /x  to  13  yit  in  diameter  occurred  in  the 
mucous  membrane  of  the  gut  and  in  the  faeces  of  a  case  of  enteritis. 


Doubtful  Species. 

In  literature  many  other  statements  are  found  as  to  the  occurrence  of 
Coccidia-like  organisms  in  different  diseases  of  man.  In  some  of  the  cases  the 
parasites  proved  to  be  fungi.  This  was  the  case  with  the  parasites  of  a  severe 
skin  disease  of  man,  formerly  called  Coccidioides  immitis  and  Coccidioides  pyogenes. 
Other  statements  are  founded  on  misapprehensions,  or  are  still  much  disputed. 
If  reference  is  here  made  to  '■'' Eimeria  hofnims"  R.  Blanchard,  1895,  this 
is  done  on  the  authority  of  the  investigator  mentioned.  The  structures  in 
question  are  nucleated  spindle-shaped  bodies  of  very  different  lengths  (18  n. 
to  100  /*),  which  either  occurred  isolated  or  were  enclosed  in  large  globular  or 
oval  cysts,  alone  or  with  a  larger  tuberculated  body  ("residual  body")-  These 
formations  were  found  by  J.  Kiinstler  and  A.  Pitres  in  the  pleural  exudation  removed 
from  a  man  by  tapping.  The  man  was  employed  on  the  ships  plying  between 
Bordeaux  and  the  Senegal  River. 

Blanchard  looks  upon  the  fusiform  bodies  as  merozoites  and  the  cysts  as 
schizonts  of  a  Coccidium.  On  the  other  hand,  Moniez  declares  the  spindle  bodies 
to  be  the  ova  and  the  supposed  residual  bodies  to  be  "  floating  ovaries "  of  an 
Echinorhynchus. 

Severi's  "monocystid  Gregarines,"  which  were  taken  from  the  lung  tissue  of  a 
still-born  child,  are  also  quite  problematical. 

No  less  doubtful  are  the  bodies  which  Perroncito  calls  Cocciditwi  jalinutn,  and 
which  he  found  in  severe  diseases  of  the  intestine  in  human  beings,  pigs,  and  guinea- 
pigs  ;  Borini  also  reported  another  case. 

'  Grunow,  "Ein  Fall  von  Protozoen  (Coccidien?)  Erkrankung  des  Darmes,"  Arch.  f. 
exper.  Path,  und  Pharm.,  1901,  xlv,  p.  262. 


H^MOSPORIDIA  151 

Order.     Haemosporidia,  Danilewsky  emend.  Schaudinn. 

The  Haemosporidia  are  a  group  of  blood  parasites,  comprising 
forms  differing  greatly  among  themselves.  Some  of  the  forms  need 
much  further  investigation.  However,  there  are  certain  true  Haemo- 
sporidia which  present  close  affinities  with  the  Coccidia,  leading 
Doflein  to  use  the  term  Coccidiomorpha  for  the  two  orders  conjoined. 
The  Haemosporidia  present  the  following  general  characteristics  : — 
(i)  They  are  parasites  of  either  red  or  white  blood  corpuscles  of 
vertebrates  during  one  period  of  their  life-history. 

(2)  They  exhibit  alternation  of  generations — asexual  phases  or 
schizogony  alternating  with  sexual  phases  or  sporogony — as  do  the 
Coccidia. 

(3)  There  is  also  an  alternation  of  hosts  in  those  cases  which  have 
so  far  been  completely  investigated.  The  schizogony  occurs  in  the 
blood  or  internal  organs  of  some  vertebrate  while  the  sporogony 
occurs  in  an  invertebrate,  such  as  a  blood-sucking  arthropod  or  leech. 

(4)  Unlike  the  Coccidia,  resistant  spores  in  sporocysts  are  not 
generally  produced,  such  protective  phases  in  the  life-cycle  being 
unnecessary,  as  the  Haemosporidia  are  contained  within  either  the 
vertebrate  or  invertebrate  host  during  the  whole  of  their  life. 

The  Haemosporidia  may  be  considered  for  convenience  under 
five  main  types  : — 

(i)  The  Plasinodiiitn  or  Hcemainceba  type.  This  includes  the 
malarial  parasites  of  man  and  of  birds.  The  asexual  multiplicative 
or  schizogonic  phases  occur  inside  red  blood  corpuscles  and  are 
amoeboid.  They  produce  distinctive,  darkish  pigment  termed  melanin 
or  haemozoin.  Infected  blood  drawn  and  cooled  on  a  slide  may 
exhibit  *'  exflagellation  "  of  the  male  gametocytes,  i.e.,  the  formation 
of  filamentous  male  gametes.  The  invertebrate  host  is  a  mosquito. 
The  malarial  parasites  of  man  are  discussed  at  length  on  p.  155. 
Similar  pigmented  haemamoeboid  parasites  have  been  described  in 
antelopes,  dogs,  and  other  mammals,  and  even  reptiles. 

(2)  The  Halteriditun  type.  The  trophozoite  stage  inside  the  red 
blood  corpuscle  is  halter- shaped.  Pigment  is  produced,  especially  near 
the  ends  of  the  organism.  The  parasites  occur  in  the  blood  of  birds. 
The  invertebrate  host  of  H.  cohunhce  of  pigeons  in  Europe,  Africa, 
Brazil  and  India,  is  a  hippoboscid  fly,  belonging  to  the  gtnviS  Lynchia. 

Halteridium  parasites  are  common  in  the  blood  of  passerine  birds, 
such  as  pigeons,  finches,  stone  owls,  Java  sparrows,  parrots,  etc. 
The  Halteridium  embraces  or  grows  around  the  nucleus  of  the  host 
red  cell  without  displacing  the  nucleus.  Young  forms  and  multipli- 
cative stages  of  H.  columhce  have  been  found  in  leucocytes  in  the 
lungs  of  the  pigeon  (fig.  76,  8-12).  Male  and  female  forms  (gameto- 
cytes) are  seen  in  the  blood  (fig.  76,  ja,  jb).     The  cytoplasm  of  the 


^52 


THE   ANIMAL   PARASITES   OF   MAN 


male  gametocytes  is  pale-staining  and  the  nucleus  is  elongate,  while 
the  cytoplasm  of  the  females  is  darker  and  the  nucleus  is  smaller 
and  round.  Formation  of  male  gametes  from  male  gametocytes 
(the  so-called  process  of  '*  exflagellation ")  may  occur  on  a  slide 
of  drawn  infected  blood,  also  fertilization,  and  formation  of  the 
ookinete,  as  first  seen  by  MacCallum.  The  correct  generic  name  for 
Halteridia  is,  apparently,  Hcumoproteiis.     Wasielewski  (191 3),  working 


/2. 


Fig.  76 — Hcemoproteus  {Halteriditim)  colunibce.  Life-cycle  diagram  :  i,  2,  stages  in  red 
blood  corpuscle  of  bird;  3,  4,  gametocytes  (3a  S ,  3b  ?);  5^,  formation  of  microgametes  ; 
6,  fertilization  (in  fly's  gut);  7,  ookinete;  8-12,  stages  in  mononuclear  leucocytes  in  lungs. 
(After  Aragao.) 


on  H,  danilewskyi  (var.  falcoiiis),  in  kestrels,  finds  that  the  halter- 
idium  may  be  pathogenic  to  nestlings.  The  cycle  of  H.  noctucv 
described  by  Schaudinn  (1904)  lacks  confirmation.  The  account  of 
the  life-cycle  of  H.  colurnhce  given  by  Aragao  (1908)  is  illustrated  in 
fig.  76.  It  agrees  with  the  work  of  Sergent  (1906-7)  and  Gonder 
(1915).  Mrs.  Adie  (19 15)  states  that  the  cycle  in  Lynchia  is  like  that 
of  a  Plasmodium. 

(3)  The  Leucocytozoon  type.     The   trophozoites  and   gametocytes 
occur  within  mononuclear  leucocytes  and  young  red  cells  (erythro- 


H^MOSPORIDIA 


153 


blasts)  in  the  blood  of  birds.  Laveran  and  Franga  consider  that 
the  Leucocytozoa  occur  in  erythrocytes.  The  host  cells  are  often 
greatly  altered  by  the  parasites,  becoming  hypertrophied  and  the  ends 
usually  drawn  into  horn-like  processes  (fig.  77),  though  some  remain 
rounded.  Leucocytozoa  are  limited  to  birds,  and  very  rarely  produce 
pigment.  Male  and  female  forms  (gametocytes)  are  distinguishable 
in  the  blood  (hg.  77),  and  the  formation  of  male  gametes  (^^ex- 
fiagellation ")  may  occur  in  drawn  blood. 

The  Leucocytozoa  were  first  seen  by  Danilewsky  in  1884.  They 
are  usually  oval  or  spherical.  It  is  not  easy  sometimes  to  distinguish 
the  altered  host  cell  from  the  parasite,  as  the  nucleus  of  the  former 

is  pushed  to  one  side  by  the 
leucocytozoon.  The  cyto- 
plasm of  the  female  parasite 
stains  deeply,  and  the  nucleus 
is  rather  small,  containing  a 
karyosome.  In  the  male  the 
cytoplasm  stains  lightly  and 
the  nucleus  is  larger,  with  a 
loose,  granular  structure. 

Many  species  of  Leuco- 
cytozoa are  recorded,  but 
schizogony  has  only  been 
described  by  Fantham  (1910)^ 
in  L.  lovati  in  the  spleen  of 
the  grouse  {Lagoptis  scoticus), 
and  by  Moldovan^  (191 3)  ^^ 
L.  ziejuanni  in  the  internal 
organs  of  screech-owls. 

M.  and  A.  Leger^  (19 14) 
propose    to    classify    Leuco- 
cytozoa, provisionally,  according  as  the   host   cells   are  fusiform  or 
rounded. 

(4)  The  Hcemogregarina  type.  Included  herein  are  many  parasites 
of  red  blood  corpuscles,  with  a  few  (the  leucocytogregarines)  parasitic 
in  the  white  cells  of  certain  mammals  and  a  few  birds.  They  are  not 
amoeboid  but  gregarine-like,  vermicular  or  sausage-shaped  (figc  7cS). 
They  do  not  produce  pigment.  They  are  widely  distributed  among 
the  vertebrata,  but  are  most  numerous  in  cold-blooded  vertebrates 
(fishes,  amphibia  and  reptiles).  The  haemogregarines  of  aquatic  hosts 
are  transmitted  by  leeches,  those  of  terrestrial  hosts  by  arthropods. 


Fig.  77. — Leucocytozoon  lovati.  a,  Male  parasite 
(microgametocyte),  within  host  cell,  whose  ends  are 
drawn  out  ;  b,  female  parasite  (macrogametocyte) 
from  blood  of  grouse.      X  1,800.     (After  Fantham.) 


'  Annals  Trop.  Med.  and  Farasitol.,  iv,  p.  255. 
2  Centralbl.f.  Bakt.,  Orig.,  Ixxi,  p.  66. 
^  Bull.  Soc.  Path.  Exot.,  vii,  p.  437. 


54 


THE   ANIMAL   PARASITES   OF   MAN 


The  nucleus  of  haemogregarines  is  usually  near  the  middle  of  the 
parasite,  but  may  be  situated  nearer  one  end.  The  body  of  the 
parasite  may  be  lodged  in  a  capsule  ('' cytocyst ").  There  is  much 
variation  in  size  and  appearance  among  haemogregarines.  Some  are 
small  {Laiikesterella) ;  some  attack  the  nucleus  of  the  host  cell  {Karyo- 
lysiis) ;  others  have  full  grown  vermicules  larger  than  the  containing 
host  corpuscle,  and  so  the  haemogregarines  bend  on  themselves  in 
the  form  of  U  (fig.  7^,  h).  Schizogony  often  occurs  in  the  internal 
organs  of  the  host,  sometimes  in  the  circulating  blood. 

The  haemogregarines  occurring  in  the  white  cells  (mononuclears 
or  polymorphonuclears)  of  mammals  have  been  referred  to  a  separate 
genus,  Lencocytogregarina  (Porter)  or  Hepatozoon  (Miller).  Such 
leucocytogregarines  are  known  in  the  dog  (fig.  79),  rat,  mouse,  palm- 
squirrel,  rabbit,  cat,  etc.  Schizogony  of  these  forms  occurs  in  the 
internal  organs,  such  as  tfie  liver,  lung  and  bone-marrow  of  the  hosts. 


Fig.  78. — Haemogregarines  from  lizards,  a,  H.  schaudinni,  var.  africana,  from  Lacerta 
ocellata ;  b,  H.  nohrei  fiom  Lacerta  muralis ;  c,  H.  marceaui  in  cytocyst,  from  Lacerta 
muralis.     (After  Fran9a.) 


They  are  apparently  transmitted  by  ectoparasitic  arthropods,  such  as 
ticks,  mites,  and  lice. 

A  few  haemogregarines  are  known  to  be  parasitic  in  the  red  blood 
corpuscles  of  mammals.  Such  are  H.  gerbilli  in  the  Indian  field  rat,. 
Gerhilliis  indicus ;  H.  balfoiiri  (jaciili)  in  the  jerboa,  Jacuhis  jacnlns, 
and  a  few  species  briefly  described  from  marsupials.  These  parasites 
do  not  form  pigment. 

Strict  leucocytic  gregarines  have  been  described  from  a  few  birds 
by  Aragao  and  by  Todd. 

The  sporogony  of  haemogregarines  is  only  known  in  a  few  cases, 
and  in  those  affinity  with  the  Coccidia  is  exhibited.  In  fact,  the 
Haemogregarines  are  now  classified  by  some  authors  with  the 
Coccidia. 

(5)  The  Babesia  or  Piroplasma  type.  These  are  small  parasites  of 
red  blood  corpuscles  of  mammals.  They  do  not  produce  pigment. 
They  are  pear-shaped,  round    or   amoeboid   in   Babesia,   bacilliform 


H^MOSPOKIDIA 


155 


and  oval  in  other  forms  referred  to  this  group.  Piroplasms  are 
transmitted  by  ticks.  These  parasites  are  described  at  length  on 
p.  172. 


Fig.  79. — Leucocytop-egarina  canis.  Life-cycle  diagram.  Constructed  from  drawings  by- 
Christophers.  (After  Castellani  and  Chalmers.)  Schizogony  occurs  in  the  bone-marrow.  The 
parasite  is  transmitted  from  dog  to  dog  by  the  tick,  Rhipicephalus  sanguineus,  development 
in  which,  so  far  as  known,  is  shown  on  the  right. 


THE    MALARIAL    PARASITES   OF    MAN. 

Malaria,  otherwise  known  as  febris  intermittens,  chill-fever,  ague, 
marsh  fever,  paludism,  etc.,  is  the  name  given  to  a  disease  of  man, 
which  begins  with  fever.  It  has  been  known  since  ancient  times  and 
is  distributed  over  almost  all  the  world,  although  very  unevenly,  but 
does  not  occur  in  waterless  deserts  and  the  Polar  regions.  In  many 
places,  especially  in  the  civilized  countries  of  Central  Europe,  the 
disease  is  extinct  or  occurs  only  sporadically,  and  large  tracts  of  land 
have  become  free  from  malaria. 

The  rhythmical  course  of  the  fever  is  characteristic.  It  begins 
apparently  suddenly  with  chilliness  or  typical  shivering,  whilst  the 
temperature  of  the  body  rises,  the  pulse  becomes  low  and  tense  and 
the  number  of  beats  of  the  pulse  increases  considerably.  After  half 
to  two  hours  the  heat  stage  begins.  The  patient  himself  feels  the  rise 
of  his  temperature  (shown  by  feeling  of  heat,  dry  tongue,  headache, 


156  THE   ANIMAL   PARASITES   OF   MAN 

thirst).  The  temperature  may  reach  41°  C  or  more.  At  the  same  time 
there  is  sensitiveness  in  the  region  of  the  spleen  and  enlargement  of 
that  organ.  After  four  to  six  hours  an  improvement  takes  place,  and 
with  profuse  perspiration  the  body  temperature  falls  rapidly,  not 
often  below  normal.  After  the  attack  the  patient  feels  languid,  but 
otherwise  well  until  certain  prodromal  symptoms  (heaviness  in  the 
body,  headache)  which  were  not  noticed  at  first,  denote  the  approach 
of  another  attack  of  fever,  which  proceeds  in  the  same  way. 

The  intervals  between  the  attacks  are  of  varying  length  which 
permit  of  a  distinction  in  the  kinds  of  fever.  If  the  attacks  intermit 
one  day,  occurring  on  the  first,  third  and  fifth  days  of  the  illness 
and  always  at  the  same  time  of  day,  it  is  termed  febris  tertiana  ;  if 
two  days  occur  between  fever  days,  it  is  called  febris  quartana.  In 
the  case  of  the  fever  recurring  daily,  later  writers  speak  of  typical 
febris  quotidiana.  But  a  quotidian  fever  may  arise  when  two  tertian 
fevers  differing  by  about  twenty-four  hours  exist  at  the  same  time 
(febris  tertiana  duplex).  The  patient  has  a  daily  attack,  but  the  fever 
of  the  first,  third  and  fifth  days  differs  in  some  point  (hour  of 
occurrence,  height  of  temperature,  duration  of  cold  or  hot  stage) 
from  the  fever  of  the  second,  fourth  and  sixth  days.  Similarly, 
two  or  three  quartan  fevers  which  differ  by  about  twenty-four  hours 
each  may  be  observed  together  {febris  quartana  duplex  or  triplex) ; 
in  the  latter  case  the  result  is  also  a  quotidian  fever. 

Two  kinds  of  tertian  fever  are  dilfefentiated — a  milder  form 
occurring  especially  in  the  spring  (spring  tertian  fever),  and  a  more 
severe  form  appearing  in  the  summer  and  autumn  in  warmer 
districts,  especially  in  the  tropics  {suninier  or  autumn  fever,  febris 
cestivo-autumnaUSy  febris  tropica,  febris  perniciosa).  The  latter  often 
becomes  a  quotidian  fever. 

All  the  afore-mentioned  infections  are  termed  acute.  They'  are 
distinguished  from  the  very  different  chronic  malarial  infection  by  the 
frequent  occurrence  of  relapses,  which  finally  lead  to  changes  of 
some  organs  and  particularly  of  the  blood.  The  relapses  are  then 
generally  marked  by  an  irregular  course  of  fever. 

The  term  masked  malaria  is  used  when  any  disturbance  of  the  state 
of  health  of  a  periodic  character  shows  itself  and  disappears  after 
treatment  with  quinine.^     Generally  it  is  a  question  of  neuralgia. 

That  intermittent  fever  was  an  infectious  disease,  although  not  one  which  was 
transmitted  direct  from  man  to  man,  had  been  assumed  for  a  long  time.  Therefore 
it  was  natural,  at  a  time  when  bacteriology  was  triumphing,  to  look  for  a  living  agent 
causmg  infection  in  malaria,  which  search  was,  seemingly,  successful  (Klebs, 
Tomasi-Crudeli,   1879).      Hence  it  was   not   surprising  that    the   discovery   of   the 

1  Quinine  is  still  almost  exclusively  the  remedy  used  in  the  treatment  of  malaria.  It  is 
prepared  from  the  bark  of  the  cinchona  tree.  This  important  remedy  was  introduced  into 
Europe  in  1640  from  Ecuador  by  Juan  del  Vego,  physician  of  ihe  Countess  del  Cinchon. 


THE    MALARIAL   PARASITES   OF   MAN  I  5/ 

real  malarial  parasites  in  November,  1880,  by  the  military  doctor  A.  Laveran^  in 
Constantine  (Algeria),  at  first  met  with  violent  opposition,  even  after  Richard 
(1882)  had  confirmed  it  and  Marchiafava,  Celli,  Grassi  and  others,  had 
further  extended  it.  Not  that  the  existence  of  structures  found  in  the  blood  of 
malaria  patients  by  Laveran  and  Richard  was  denied  ;  on  the  contrary,  the  investiga- 
tions of  the  opponents  furnished  many  valuable  discoveries,  but  the  organisms  were 
differently  interpreted  and  considered  to  be  degeneration  products  of  red  blood  cor- 
puscles. Only  when  Marchiafava  and  Celli  (1885)  saw  movements  in  the  parasites, 
wiiich  Laveran  called  Oscillaria  malarice  and  later  Hcematozoon  malarice^  was  their 
animal  nature  admitted  and  the  parasites  were  r\7kx^^di  Plasmodium  malaricE.  Shortly 
before  this,  Gerhardt  (1884)  had  stated  that  the  disease  could  be  transmitted  by  the 
injection  of  the  blood  of  a  malarial  patient  to  a  healthy  person. 

This  supplied  the  starting  point  for  further  investigations,  which  were  made  not 
exclusively,  but  principally,  by  Italian  investigators  (Golgi,  Marchiafava  and  Celli, 
Bignami  and  Bastianelli,  Grassi  and  Feletti,  Mannaberg,  Romanowsky,  Osier, 
Thayer  and  others).  In  1885  Golgi  described  the  asexual  cycle  in  the  blood, 
in  the  case  of  the  quartan  parasite.  These  investigations,  after  attention  had  been 
drawn  by  Danilewsky  (1890)  to  the  occurrence  of  similar  endoglobular  parasites  in 
birds,  were  extended  to  the  latter  (Grassi  and  P'eletti,  Celli  and  Sanfelice^Kruse, 
Labbe  and  others). 

The  result  was  as  follows  :  Malaria  in  man  (and  birds)  is  the  result  of  peculiar 
parasites  included  in  the  Sporozoa  by  Metchnikoff,  which  parasites  live  in  the  erythro- 
cytes, grow  in  size  and  finally  "  sporulate,"  that  is,  separate  into  a  number  of  "  spores  " 
which  leave  the  erythrocytes  and  infect  other  blood  corpuscles.  Morphologically  and 
biologically  several  species  (and  respectively  several  varieties)  of  malarial  parasites 
may  be  distinguished,  on  which  the  different  intermittent  forms  depend.  Trans- 
mission of  the  blood  of  patients  to  healthy  people  produces  a  malarial  affection  which 
corresponds  in  character  to  the  fever  of  the  patient  from  whom  the  inoculation  was 
made.  The  combined  types  of  fever  (tertiana  duplex,  quartana  duplex  or  triplex) 
are  explained  by  the  fact  that  the  patient  harbours  two  or  three  groups  of  parasites 
which  differ  in  their  development  by  about  twenty-four  hours,  whilst  the  irregular  fevers 
depend  on  deviation  from  the  typical  course  of  development  of  the  parasites.  In 
addition  to  stages  of  the  parasites  which  could  easily  be  arranged  in  a  developmental 
series  concurrent  with  the  course  of  the  disease,  other  phases  of  the  parasites  also 
became  known,  such  as  spheres,  crescents,  polymitus  forms,  which  seemed  not  to  be 
included  in  the  series  and,  therefore,  were  very  differently  interpreted. 

The  decision  reached  at  the  beginning  of  the  last  decade  of  the  last  century,  which 
found  expression  in  comprehensive  statements  (Mannaberg,  Ziemann  and  others), 
only  concerned  a  part  of  the  complete  development  of  the  malarial  parasites.  No  one 
could  with  any  degree  of  certainty  demonstrate  how  man  became  infected,  nor  were 
there  reliable  hypotheses  based  on  analogy  with  other  parasites  concerning  the  exit 
of  the  excitants  of  malaria  from  the  infected  person  and  their  further  behaviour. 
Numerous   hypotheses    had   been    advanced,  but   none  was    able    to    elucidate  the 

'  The  discovery  of  Laveran  is  in  no  way  lessened  by  the  fact  that  one  investigator  or 
another  (according  to  Blanchard  {^Arch.  de  Paras.,  vii,  1903,  p.  152],  P.  F.  H.  Klencke 
in  1843)  had  seen,  mentioned  and  depicted  malarial  parasites.  {Neiie  phys.  Abhandl, 
auf.  selbstdnd.  Beob.  gegr.,  Leipzig,  1843,  p.  163,  fig.  25).  In  1847  Meckel  had  recognized 
that  the  dark  colour  of  the  organs  in  persons  dead  of  malaria  was  due  to  pigment.  Virchow 
in  1848  stated  that  this  pigment  occurred  in  blood  cells.  Kelsch  in  1875  recognized  the 
frequency  of  melaniferous  leucocytes  in  the  blood  of  malarial  patients.  Beauperthuy  (1853) 
noticed  that  in  Guadeloupe  there  was  no  malaria  at  altitudes  where  there  were  no  '*  insectes 
tipulaires,"   and  suggested  that  the  disease  was  inoculated  by  insects. 


158  THE   ANIMAL   PARASITES   OF   MAN 

various  observations  made  from  time  to  time  in  dealing  with  malaria.  One  hypo- 
thesis only  seemed  to  have  a  better  foundation.  Manson  (1894),  who  knew  from  his 
own  experience  the  part  played  by  mosquitoes  in  the  development  of  Filaria  from 
the  blood  of  man,  applied  this  also  to  the  malarial  parasites  living  in  the  blood, 
whereby  at  least  the  way  was  indicated  by  which  the  Haemosporidia  could  leave  man. 
The  parasites  were  said  finally  to  get  into  water  through  mosquitoes  which  had 
sucked  the  blood  of  malarial  patients,  and  the  germ  spread  thence  to  men  who  drank 
the  water.  In  some  cases  the  parasites  were  supposed  to  reach  man  by  the  inhaling  of 
the  dust  of  dried  marshes.  On  the  other  hand,  Bignami  believed  that  the  mosquitoes 
were  infected  in  the  open  air  by  malarial  parasites  which  occurred  there  in  an  unknown 
stage  and  the  insects  transmitted  the  germs  to  man  when  biting.  R.  Koch  combined 
both  hypotheses,  without,  however,  producing  positive  proof.  R.  Ross,  then  (1897-8) 
an  English  military  doctor  in  India,  was  the  first  to  succeed  in  this.  He  had  been 
encouraged  by  Manson  to  study  the  fate  of  malarial  Plasmodia  which  had  entered 
the  intestine  of  mosquitoes  with  malaria-infected  blood,  especially  in  the  case  of  the 
Plasmodium  {Proteosojna)  living  in  the  blood  of  birds.  He  showed  that  the  Proteo- 
soma  penetrate  the  intestmal  wall  of  the  mosquitoes,  grow  and  develop  into  large 
cysts  which  produce  innumerable  rod-like  germs,  which  burst  into  the  body  cavity 
and  penetrate  the  salivary  glands.  Ross  allowed  mosquitoes  to  suck  the  blood  of 
birds  affected  by  malaria,  and  some  nine  days  later,  let  the  infected  mosquitoes 
which  had  been  isolated  suck  healthy  birds.  After  five  to  nine  days  Proteosoma 
were  found  to  occur  in  the  blood  of  the  birds  used.  The  Proteosoma  and  Halteridium 
of  birds  were  also  further  investigated  by  MacCallum  (1897-8),  Koch  and  others,  and 
important  results  followed. 

In  any  case  Ross  (1898)  had  clearly  established  the  importance  of  mosquitoes  in 
the  spread  of  malaria  among  birds.  It  was  now  only  a  question  of  proving  whether, 
and  how  far,  mosquitoes  were  concerned  with  human  malaria.  Ross  himself  worked 
to  this  end.  Here  the  experiments  of  Italian  investigators  (Bignami,  Bastianelli, 
Grassi)^  were  of  importance.  These  investigators  studied  the  fate  of  malarial 
parasites  in  man,  produced  malaria  in  men  experimentally  by  the  bites  of  infected 
mosquitoes,  and  established  that  only  mosquitoes  belonging  to  the  genus  Anopheles 
were  concerned,  and  not  species  of  Culex.  These  laiter  are  only  able  to  transmit 
Proteoso7na  to  birds.  It  is  true  that  Culex  can  ingest  the  human  malarial  parasites, 
but  the  latter  do  not  develop  in  them.  Development  only  occurs  in  species  of 
Anopheles.  In  Anopheles  (and  similarly  for  Proteosoma  in  Culex)  sexual  repro- 
duction takes  place  ;  crescents,  spheres  and  polymitus  forms  are  necessary  stages 
of  development  in  the  mosquito. 

With  these  discoveries  the  campaign  against  malaria  became  more  definite.  It 
was  directed  partly  against  the  transmitters,  whose  biology  and  life-cycle  were  more 
thoroughly  investigated,  instead  of  merely  against  the  infection  of  the  adult  Anopheles. 
The  latter  do  not,  as  was  believed  for  some  time,  transmit  the  malarial  germs  to  their 
offspring.  They  always  infect  themselves  from  human  beings,  whereby  the  relapses 
appearing  in  early  summer,  and  the  latent  infection,  especially  of  children  of  natives, 
play  a  principal  part  (Stephens  and  Christophers,  Koch).  Further,  the  crusade  was 
directed  against  the  infection  of  man  by  the  bites  of  Anopheles.  Important  results 
have  been  obtained  in  these  directions.  Low  and  Sambon  in  1900  lived  in  a 
mosquito-screened  hut  in  a  malarial  part  of  the  Roman  Campagna  for  three  of  the 
most  malarious  months  and  did  not  contract  the  disease.  In  the  same  year  Dr.  P.  T. 
Manson  was  infected  with  malaria  by  infected  mosquitoes  sent  from  Italy.  The  role 
of  mosquitoes  having  been  proved,  it  may  be  hoped  that  ultimately  the  eradication 
of  malaria,  or  at  least  a  considerable  restriction  of  it,  will  be  achieved. 

*  Grassi,  B.  (1901),  "  Die  Malaria,"  250  pp.,  8  plates.     G-  Fischer,  Jena. 


DEVELOPMENT   OF   THE    MALARIAL    PARASITES   OF   MAN  1 59 

It  is  of  importance  to  record  that,  although  malarial  parasites  occur  in  mammals 
(monkeys,  bats,  etc.)  the  human  ones  are  not  transmissible  to  mammals,  not  even  to 
monkeys.  The  species,  therefore,  are  specific  to  the  different  hosts  (Dionisi,  Kossel, 
Ziemann,  Vassall). 

An  important  work  dealing  with  the  modern  applications  of  the  mosquito-malaria 
theory  in  all  parts  of  the  Tropics  was  published  by  Sir  Ronald  Ross  in  191 1.  It  is 
entitled  "The  Prevention  of  Malaria"  (John  Murray,  London,  21s.). 


DEVELOPMENT  OF  THE  MALARIAL  PARASITES  OF  MAN. 

The  commencement  of  the  developmental  cycle  and  of  the  infection 
of  man,  is  the  sporozoites  (fig.  80,  /)  which  are  passed  into  the  blood 
of  a  person  by  the  bite  of  an  infected  mosquito.  Prior  to  this  the  para- 
sites collect  in  the  excretory  ducts  of  the  salivary  glands  (fig.  80,  27) 
of  the  Anopheles.  The  sporozoites  are  elongate  and  spindle-shaped, 
10  fx  to  20  fi  long  and  1  //,  to  2  //,  broad,  v^'ith  an  oval  nucleus  situated 
in  the  middle.  They  are  able  to  glide,  perform  peristaltic  contractions, 
or  curve  laterally.  Schaudinn  has  studied  the  penetration  of  the  red 
blood  corpuscles  (fig.  80,  2)  by  the  sporozoites  in  the  case  of  the 
living  tertian  parasite.  The  process  takes  forty  to  sixty  minutes  in 
drawn  blood.  After  its  entrance  the  parasite,  which  is  now  called  a 
trophozoite,  contracts,  and  becomes  an  active  amoebula  (fig.  80,  5). 
It  develops  a  food  vacuole  and  grows  at  the  expense  of  the  invaded 
blood  corpuscle  (fig.  80,  4),  which  is  shown  by  the  appearance  of 
pigment  granules  (transformed  haemoglobin)  in  it.  When  the  maxi- 
mum size  is  attained,  multiplication  by  schizogony  (fig.  80,  ^-y)  begins 
with  a  division  of  the  nucleus,  which  is  followed  by  further  divisions  of 
the  daughter  nuclei,  the  number  of  which  varies  up  to  16  or  even  32, 
depending  on  the  species  of  the  parasite.  Then  the  cytoplasm  divides 
into  as  many  portions  as  there  are  nuclei,  the  result  being  a  structure 
suggestive  of  the  spokes  of  a  wheel  or  of  a  daisy,  the  centre  of  the 
resulting  rosette  being  occupied  by  dark  pigment.  Finally,  the  parts 
separate  from  one  another,  leaving  behind  a  residual  body  contain- 
ing the  pigment,  and  the  daughter  forms  issue  into  the  blood  plasma 
as  merozoites  (fig.  80,  7).  They  are  actively  amoeboid  (fig.  80,  8)  and 
soon  begin  to  enter  other  blood  corpuscles  of  their  host,  for  the  entry 
into  which  thirty  to  sixty  minutes  are  necessary,  according  to 
Schaudinn's  observations.  ^ 

Here  they  behave  like  sporozoites  which  previously  entered  and 

'  It  should  be  remembered  that  some  authors  (Laveran,  Argutinsky,  Panichi,  Serra)  argue 
against  the  intra-globular  position  of  malarial  parasites  and  state  that  they  only  adhere  out- 
wardly to  the  red  blood  corpuscles.  These  views  have  recently  been  revived  by  Mary  Rowley- 
Lawson,  and  she  states  that  the  malarial  parasite  is  "  extracellular  throughout  its  life-cycle  and 
migrates  from  red  corpuscle  to  red  corpuscle  destroying  each  before  it  abandons  it."  {/ourn. 
Exper.  Med.^  1914,  xix,  p.  531.) 


6 


3 


ff)"^^fi'  "D^'^^'-wf 


7 

V 


Fig.  8o. — Life-cycle  of  the  tertian  parasite  [Plasmodium  vivax).  Figs,  i  to  17,  X  1,200; 
figs.  18  to  27,  X  600.  (After  Llihe,  based  on  figures  by  Schaudinn  and  Grassi.)  i,  sporozoite  ; 
2,  entrance  of  the  sporozoite  into  a  red  blood  corpuscle  ;  3,  4,  growth  of  the  parasite,  now 


DEVELOPMENT   OF   THE   MALARIAL   PARASITES   OF   MAN  l6l 

again  produce  merozoites.  This  process  is  repeated  until  the  number 
of  parasites  is  so  large  that,  at  the  next  migration  of  the  merozoites,  the 
body  of  the  person  infected  reacts  with  an  attack  of  fever/  which  is 
repeated  with  the  occurrence  of  the  next  or  following  generations. 

The  growth  and  schizogony  last  different  times,  according  to  the 
species  of  the  parasite,  about  forty-eight  hours  in  the  case  of  the 
parasite  of  febris  tertiana  or  tropica,  and  seventy-two  hours  for  the 
quartan  parasite.  The  various  intermittent  forms  produced  by  them 
depend  on  this  specific  difference  in  the  malarial  parasites. 

The  schizogony  can,  however,  only  be  repeated  a  certain  number 
of  times,  supposing  that  the  disease  has  not  been  checked  prematurely 
by  the  administration  of  quinine,  which  is  able  to  kill  the  parasites. 
It  appears  that  after  a  number  of  attacks  of  fever  the  conditions  of 
existence  in  man  are  unfavourable  for  the  malarial  parasites,  and  this 
brings  about  the  production  of  other  forms  which  have  long  been 
known,  but  also  long  misunderstood  (spheres,  crescents,  polymitus). 
The  merozoites  in  this  case  no  longer  grow  into  schizonts,  or  at  least 
not  all  of  them,  but  become  sexual  individuals  called  gametocytes 
(fig.  80,  9 — 12),  which  only  start  their  further  development  when  they 
have  reached  the  intestine  of  Anopheles.  This  does  not  take  place 
in  every  case,  nor  with  all  the  gametocytes  which  exist  in  the  blood  of 
patients  with  intermittent  fever.  Of  those  parasites  which  remain 
in  the  human  blood  the  male  ones  (microgametocytes)  soon  perish, 
the  females  (macrogametocytes)  persist  for  some  long  time,  and  per- 
haps at  last  acquire  the  capacity  of  increasing  by  schizogony.  They 
might  thus  form  merozoites  which  behave  in  the  body  as  if  they  had 
proceeded  from  ordinary  schizonts  (fig.  80,  13c — i^c).  If  their 
number  increases  sufficiently,  in  course  of  time  the  patient,  who 
was  apparently  recovering,  has  a  new  series  of  fever  attacks,  or 
relapses,  without  there  having  been  a  new  infection.  This  is  the 
view  of  Schaudinn,  who  from  researches  of  his  own  concluded  that 
relapses  were  brought  about  by  a  sort  of  parthenogenetic  reproduction 
of  macrogametocytes.     R.  Ross,  on  the  contrary,  believes  that  in  the 

sometimes  called  a  trophozoite  ;  5,  6,  nuclear  division  in  schizont ;  7,  free  merozoites  ;  8,  the 
merozoites  which  have  developed  making  their  way  into  blood  corpuscles,  (arrow  pointing  to  the 
left)  and  increase  by  schizogony  (3 — 7) ;  after  some  duration  of  disease  the  sexual  individuals 
appear ;  ga — 12a,  macrogametocytes  ;  gd — 12(5,  microgametocytes,  both  still  in  the  blood-vessels 
of  man.  If  macrogametocytes  (12a)  do  not  get  into  the  intestine  of  Anopheles  they  may  perhaps 
increase  parthenogenetically  according  to  Schaudinn  (12a;  13^ — 17^).  The  merozoites  which 
have  arisen  (I7(:)  become  schizonts  3 — 7.  The  phases  shown  underneath  th^  dotted  line 
(13 — 17)  proceed  in  the  stomach  of  Anopheles.  I'^^b  and  14/^,  formation  of  microgametes ;  13a 
and  14a,  maturation  of  the  macrogametes ;  15*^,  microgamete  ;  16,  fertilization  ;  17,  ookinete; 
18,  ookinete  in  the  walls  of  the  stomach  ;  19,  penetration  of  the  epithelium  of  the  stomach  ; 
20 — 25,  stages  of  sporogony  on  the  outer  surface  of  the  intestinal  wall ;  26,  migration  of  the 
sporozoites  to  the  salivary  gland  ;  27,  salivary  gland  with  sporozoites. 

*  The  incubation  period,  that  is,  the  time  between  infection  and  the  first  attack  of  fever,  is 
ten  to  fourteen  days  ;  with  severe  infection  fewer  days  (minimum  5  to  6)  are  needed. 

II 


l62  THE   ANIMAL   PARASITES   OF   MAN 

relatively  healthy  periods  the  number  of  parasites  in  the  blood  falls 
below  that  necessary  to  provoke  febrile  symptoms ;  relapses  then 
result  merely  from  increase  in  the  numbers  of  the  parasites  present  in 
the  individual.     Ross's  view  is  now  generally  accepted. 

If   the   gametocytes,   which    are   globular,    or   in    the    pernicious 
or  malignant  tertian  parasite  crescentic  (fig.    8i),  gain  access  to  the 

"  I  ©  (J  ©  <7^ 

a  h  c  d  e 

Fig.  8i. — Stages  of  development  of  pernicious  or  malignant  tertian  parasites  in  the  intestine 
of  Anopheles  maculipennis.  (After  Grassi.)  a,  macrogametocyte  (crescent)  still  attached  to 
human  blood  corpuscles;  b,  macrogametocyte  (sphere)  half  an  hour  after  ingestion  by  the 
mosquito ;  <r,  microgametocyte  (crescent)  attached  to  the  blood  corpuscle  ;  d,  microgameto- 
cyte  (sphere)  half  an  hour  after  ingestion  ;  the  nucleus  has  divided  several  times ;  ^,  micro- 
gametes  attached  to  the  residual  body  (polymitus  stage). 

intestine  of  an  Anopheline/  they  mature.  The  macrogametocytes 
extrude  a  part  of  their  nuclear  substance  (fig.  80,  i^a,  14a)  and 
thereby  become  females  or  macrogametes.  The  microgametocytes,  on 
the  other  hand,  undergo  repeated  nuclear  division,  preparation  for 
this  being  made  apparently  whilst  in  the  blood  of  man.     This  results 


Fig.  82. — Ookinete  of  the  malignant  tertian  parasite  in  the  stomach  oi  Anopheles  maculipennis ^ 
thirty-two  hours  after  ingestion  of  blood.     (After  Grassi.) 

in  the  formation  of  threadlike  bodies  which  move  like  flagella  and 
finally  detach  themselves  from  the  residual  body  (fig.  80,  736,  i^h). 
These  are  the  males  or  microgametes^  (fig.  80,  15^)- 

Copulation  takes  place  in  the  stomach  of  the  Anopheline  (fig.  80, 16). 
A  microgamete  penetrates  a  macrogamete  and  coalesces  with  it.     The 

*  Schizonts  ingested  about  the  same  time  perish  in  the  intestine  of  the  mosquito. 

2  If  the  microgametocytes  are  sufficiently  mature  the  formation  of  microgametes  occurs  in 
the  blood  of  man  as  soon  as  it  is  taken  from  the  blood-vessel  and  has  been  cooled  and 
diluted.  Such  a  stage  is  called  a  Polymihts  form,  and  the  process  has  been  called 
'•  exflagellation." 


DEVELOPMENT   OF   THE    MALARIAL   PARASITES   OF   MAN 


163 


Fig.  83.  —  Section  of  the 
stomach  of  an  Anopheles^  with 
cysts  (oocysts)  of  the  malignant 
tertian  parasite.     (After  Grassi). 


fertilized   females   elongate    very   soon    and    are    called   ookinetes  or 

"  vermicules "  (figs.    80,    ij ;  82).     They   penetrate    the   walls  of  the 

stomach,  pierce    the  epithelium  (fig.  80,  18,   19),  and   remain    lying 

between     it    and     the    superficial     stratum 

(tunica    elastico  -  muscularis).      Then    they 

become    rounded    and    gradually    develop 

into     cysts    which     grow    larger    and    are 

finally    visible     to     the    naked    eye,    being 

called  oocysts  (figs.  80,  20-24. ;   83).     Their 

size   at    the    beginning    is   about    5   fx,    the 

maximum    that    they    attain    is    60   //,,    only 

exceptionally  are  they  larger. 

The  sporulation  (figs.  80,  21 — 25;  84), 
which  now  follows,  begins  with  repeated 
multiple  fission  of  the  nucleus.  Long 
before  the  definitive  number  of  nuclei, 
which  varies  with  the  individual,  is  attained 

the  protoplasm,  according  to  Grassi,  begins  to  segment  around  the 
individual  large  nuclei  but  without  separating  completely  into  cell 
areas.  According  to  Schaudinn, 
however,  there  is  a  condensation 
of  the  outstanding  protoplasmic 
strands.  It  is  certain  that  the  num- 
ber of  nuclei  increases  with  simul- 
taneous decrease  in  size.  They 
soon  appear  on  the  surface  of  the 
strands  or  sporoblasts,  surround 
themselves  with  some  cytoplasm 
and  then  elongate  (fig.  84).  In 
this  manner  the  sporozoites  are 
formed  and  break  away  from  the 
unused  remains  of  the  cytoplas- 
mic strands  of  the  sporoblasts 
(fig.  80,  26).  The  number  of  the 
sporozoites  in  an  oocyst  varies  from 
several  hundreds  to  ten  thousand. 


Fig.  84. — Four  different  sporulation  stages 
of  malarial  parasites  from  Anopheles  maculi- 
pennis,  much  magnified.  a — ^,  of  the 
malignant  tertian  parasite  ;  a,  four  to  four 
and  a  half  days  after  sucking  ;  t  and  c,  five 
to  six  days  after  sucking  ;  d,  of  the  tertian 
parasite,  eight  days  after  sucking.  (After 
Grassi.) 


The  sporulation  is  influenced  in  its 
duration  by  the  external  temperature 
(Grassi,  Jansci,  Schoo).  In  the  tertian 
parasite  it  takes  place  quickest  at  a 
temperature  of  25°  to  30°  C.  and  takes 
eight  to    nine   days.      A  temperature    a 

few  degrees  lower  has  a  retarding  effect  (eighteen  to  nineteen  days  at  18°  to  20°  C). 
A  still  lower  one  has  a  restraining  or  even  destructive  effect.  Temperatures  over 
35°  C.  also  exercise  a  harmful  effect.  The  malignant  tertian  parasite  seems  to  need 
a  somewhat  higher  temperature  and  the  quartan  parasite  a  lower  one. 


l64  THE   ANIMAL   PARASITES   OF    iMAN 

The  sporozoites  of  the  various  malarial  parasites  show  no  specific 
differences.  They  were  stated  by  Schaudinn  to  occur  in  three  forms, 
and  these  were  described  as  indifferent  (neuter),  female  and  male. 
There  is,  however,  little  or  no  evidence  for  this  hypothetical  differ- 
entiation. The  last  were  said  to  perish  prematurely,  that  is,  in  the 
oocyst.  The  others  after  the  rupture  of  the  oocysts  enter  the  body 
cavity  of  the  Anophelines,  whence  they  are  carried  along  in  the  course 
of  the  blood.  Finally  they  penetrate  the  salivary  glands  (fig.  80,  2^) 
probably  by  their  own  activity,  break  through  their  epithelia  and 
accumulate  in  the  salivary  duct  (fig.  80,  27).  At  the  next  bite  by  the 
mosquito  they  are  transmitted  to  the  blood-vessels  of  man. 

The  Species  oy  Malarial  Parasites  of  Man. 

In  view  of  the  differences  in  opinion  regarding  "species"  and  "varieties,"  the  dispute 
whether  the  malarial  parasites  of  man  represent  one  species  with  several  varieties,  or  several 
species  is  almost  superfluous.     If  necessary  two  genera  may  be  distinguished. 

The  parasites  of  the  tertian  and  quartan  fever  are  alike  in  that 
their  gametocytes  have  a  rounded  shape  (figs.  80,  /2,  13),  whilst  the 
corresponding  stages  of  the  pernicious  or  malignant  tertian  parasites 
are  crescentic  (figs.  81,  88).  These  differences  are  used  by  some  writers 
as  the  distinguishing  characteristic  of  two  genera  :  Plasmodium^  Mar- 
chiafava  and  Celli,  1885,  for  the  first  mentioned  species  ;  Laverania^ 
Grassi  and  Feletti,  1889,  for  the  pernicious  or  malignant  tertian  parasite. 
Whether  there  is  a  genuine  quotidian  fever  and  accordingly  a  special 
quotidian  parasite  is  still  disputed. 

These  parasites  are  treated  in  practical  detail  in  Stephens  and  Christophers' 
"  Practical  Study  of  Malaria,"  3rd  edition,  1908. 

Plasmodium  vivax,  Grassi  and  Feletti,   1890. 

Syn.  :  Hcsmamceba  vivax,  Grassi  and  Feletti,  1890;  Plasmodium  malaricB  van 
tertiancE,  Celli  and  Sanfelice,  1891  ;  HcFmajiiccba  laverani  var.  teriiana,  Labbe, 
1894  ;  HcBmosporidium  tertianum,  Lewkowitz,  1897  ;  Plasmodium  malaricE  tertianum, 
Labbe,  1899:  HcEmamceba  malaria  var.  magna,  Laveran,  1900,  p.p.;  Hcemamceba 
malaricB  var.  tertiancB,  Laveran,  1901. 

This  species,  P.  vivax,^  is  the  causal  agent  of  the  simple  or  spring 
tertian  fever  and  is,  therefore,  named  directly  the  tertian  or  benign 
tertian  parasite  (figs.  80,  3-8  ;  85).  During  the  afebrile  period  in  the 
patient,  the  young  trophozoites  or  amcebulae  appear  on  or  in  the  red 
blood  corpuscles  as  pale  bodies  of  1-5  /^  to  2  //,  diameter  which  at 
first  show  only  slow  amoeboid  movements.  Their  nucleus  is  difficult 
to  recognize  in  the  early  stage.  Soon  the  food  vacuole  is  formed  and 
this  grows  concomitantly  with  the  trophozoite  and  the  parasite  has  a 
ring-like  appearance.     Afterwards  the  vacuole  diminishes,  and  at  this 


See  Schaudinn,  F.  (1902),  Arb.  a.  d.  kaiserl.  Gesundheits.,  xix,  pp.  169-250,  3  plates. 


PLASMODIUM   VIVAX 


165 


period  the  first  brownish  melanin  granule  appears.  From  this  time 
the  activity  and  number  of  the  pigment  granules  increase  with  con- 
tinuous growth.  When  the  parasite  has  grown  to  about  one-third 
the  diameter  of  the  erythrocyte  the  latter  shows  characteristic  red 
Schiiffner's  dots  or  "  fine  stippling/'  after  staining  with  Romanowsky's 
solution.  Later,  after  about  twenty-four  hours,  the  blood  corpuscles 
begin  to  grow  pale,  then  to  increase  in  size,  and  after  thirty-six  hours, 
that  is,  about  twelve  hours  before  the  next  attack  of  fever,  schizogony 
of  the  parasite  is  initiated  by  the  division  of  the  nucleus.  The  parasite 
at  this  time  occupies  half  to  two-thirds  of  the  enlarged  blood  cor- 
puscle. The  daughter  nuclei  continue  dividing  until  sixteen,  and 
occasionally  twenty-four,  daughter  nuclei  are  produced.  The  pigment 
which,  up  till  now  lies  nearer  the  periphery,  moves  to  the  middle, 
while  the  nuclei  lie  nearer  the  surface. 


Fig.  85. — Development  of  the  tertian  parasite  in  the  red  blood  corpuscles  of  man  ; 
on  the  right  a  "  Polymitus."     (After  Mannaberg.)     See  also  fig.  80,  j — 7. 

Around  each  nucleus  a  portion  of  cytoplasm  collects  and  thus 
young  merozoites  are  produced.  These  separate  from  each  other  and 
from  the  little  residual  masses  ^  which  contain  the  melanin  and  pass 
from  the  blood  corpuscles,  which  now  can  hardly  be  recognized,  to 
the  blood  plasma,  where  they  soon  attack  new  erythrocytes. 

The  migration  of  the  merozoites  initiates  a  new  attack  of  fever 
and  two  groups  of  tertian  parasites  in  the  blood,  differing  in  develop- 
ment by  about  twenty-four  hours,  are  the  conditions  for  febris  tertiana 
duplex. 

After  a  lengthy  duration  of  fever  the  gametocytes  (figs.  80,  9 — 12) 
appear.  They  are  uninucleate.  The  microgametocytes  are  about  the 
size  of  fully  developed  schizonts,  the  macrogametocytes  are  somewhat 
larger.     Their  further  development  takes  place  in  Anophelines. 


'  The  pigment  masses  (melanin  or  hsemozoin)  are  taken  up  by  the  leucocytes,  particularly 
the  mononuclear  ones,  and  are  carried  especially  to  the  spleen,  and  also  to  the  liver  and  the 
bone-marrow.  From  this  circumstance  arises  the  v^^ell-known  pigmentation  of  the  spleen  in 
persons  who  have  suffered  from  malaria. 


1 66 


THE   ANIMAL   PARASITES   OF   MAN 


The  chief  distinctive  characteristics  of  the  simple  tertian  parasite, 
as  seen  in  infected  blood,  are  :— (i)  The  infected  red-cell  is  usually 
enlarged  ;  (2)  the  presence  of  fine  red  granules  known  as  Schiiffner's 
dots  in  the  red  blood  corpuscles,  after  Romanowsky  staining;  (3) 
the  fragile  appearance  of  the  parasite  compared  with  other  species. 
Large  forms  are  pigmented,  irregular  and  *' flimsy-looking,"  some- 
times appearing  to  consist  of  separate  parts.  Irregularity  of  contour 
is  common. 

Ahmed  Emin'  (1914)  has  described  a  small  variety  of  P.  vivax. 

Plasmodium    malariae,   Laveran. 

Syn.  :  Oscillaria  malaricz,  Laveran,  p.p.,  1883  ;  Hcernamceba  malaria,  Gr.  et  Fel., 
1890;  Plasmodium  malarice  var.  quartancE^  Cell!  et  Sanfel.,  1891  ;  Hcemamoeba 
laverani  var.  quartana  Labbe,  1894  ;  Hamosporidium  quartan<z,  Lewkowitz,  1897  ; 
Plasmodium  malaria  quartanum,  Labbe,  1899  ;  P las 7110 diuin  golgii,  Sanibon,  1902  ; 
Laverania  malarice,  Jancso,  1905  nee  Grassi  et  Fel.  1890  ;  HcEinamoeba  malarice 
var.  quartance  ;    Lav.,  1901. 

Plasmodium  malarice  is  the  parasite  of  quartan  malaria  (fig.  86). 
The  trophozoites  of  the  quartan  parasite  differ  from  the  corresponding 
stages  of  the  tertian  parasite  in  that  their  motility  is  less  and  soon 
ceases.  They  differ  also  in  their  slower  growth,  by  the  early 
disappearance  of  the  food  vacuole,  by  the  more  marked  formation 
of  the  dark  brown  pigment,  and  by  the  fact  that  the  red  blood 
corpuscles  attacked  are  not  altered  either  in  colour  or  size. 


Fig.  86. — Development  of  the  quartan  parasite  in  the  red  corpuscles  of  man— asexual 
stages.     (After  Manson.) 

When  the  parasites  have  grown  almost  to  the  size  of  the  erythrocytes 
schizogony  occurs.  The  pigment  granules  arrange  themselves  in  lines 
radiating  towards  the  centre  and  the  merozoites  are  also  radially 
disposed  in  groups  of  6,  8,  10  or  even  12,  but  are  often  arranged 
less  regularly.  The  whole  development,  growth  and  schizogony, 
occupies  seventy-two  hours. 


Bull.  Soc.  Path.  Exot.,  vii,  p.  385. 


LAVERANIA   MALARIA  167 

The  appearance  of  quartana  duplex  or  triplex  is  conditional  on  the 
presence  in  the  blood  of  the  patient  of  two  or  three  groups  of 
Plasmodia  differing  in  their  development  by  twenty-four  hours. 

The  chief  distinctive  characters  of  the  quartan  parasite  are  :  (i) 
The  erythrocyte  is  unchanged  in  size  ;  (2)  the  rings  are  compact  and 
show  pigment  early;  in  the  larger  forms  the  chromatin  is  dense  and 
relatively  plentiful ;  (3)  the  pigment,  which  is  relatively  well-marked, 
may  be  arranged  at  the  periphery. 

Laverania  malarias,  Grassi  and   Feletti,  1890  =  Plasmodium 
falciparum,  Welch,   1897. 

Syn.  :  Plasmodium  malarice  var.  quotidiance^  Celli  et  Sanf.,  1891  ;  Hcemamoeba 
malaricB  prczcox;  Gr.  et  Fel.,  1892  (nee  H.  prcecox^  Gr.  et  Fel,  1890)  ;  Hcemamoeba 
laverani,  Labbe,  1894  ;  HcEmatozoon  falciparum,  Welch,  1897  ;  Hcemosporidium 
undecimancE  and  H.  sedecimancp  a?td  H.  vigesimo-tertiana,  Lewkowitz,  1897  ;  Hcsma- 
mceba  malarice  parva.  Lav.,  1900;  Plasmodium  prcecox,  Dofl.,  1901  ;  Plasmodium 
imfnaculatu7n,  Schaud.,  1902  ;  Plasmodium  falciparum,  Blanch.,  1905. 

The  names  most  commonly  used  for  the  parasite  of  malignant 
tertian  malaria  are  Plasniodliun  falcipamni  and  Laverania  malarice. 

The  summer  and  autumn  fever  (febris  aestivo-autumnalis),  also 
called  malignant  tertian  or  sub-tertian,  is  caused  by  a  malarial  parasite 
which  is  distinguished  by  the  small  size  of  its  schizont,  while  the 
gametocytes  are  crescentic  (figs.  81,  88). 

Most  authors  identify  this  kind  of  fever  or  the  parasites  which  cause  \t  {Laverania 
malarice)  with  the  pernicious  malaria  of  the  tropics.  Ziemann,  however,  repeatedly 
has  drawn  attention  to  certain  small  but  definite  differences  between  the  usual  malig- 
nant tertian  or  pernicious  parasites  which  occur  in  the  tropics  and  the  tropical  parasites 
of  some  malarial  districts,  particularly  of  West  Africa,  and  insists  that  at  least  two 
varieties  or  sub-species  occur.  Other  investigators  distinguish  from  this  or  these 
forms  a  quotidian  parasite.  On  the  other  hand,  the  assertion  is  made  that  there  are 
no  specific  differences,  but  that  the  malignant  or  pernicious  tertian  parasite  which 
normally  needs  forty-eight  hours  for  its  development  in  the  blood  of  man,  can 
also  develop  in  twenty-four  hours.  The  establishment  of  the  duration  of  the 
development  is  a  matter  of  especial  difficulty,  because  the  stages  of  schizogony 
are  far  less  numerous  in  the  peripheral  blood  than  in  that  of  the  internal  organs.  It 
is  also  stated  that  the  tropical  parasite  very  seldom  forms  crescentic  but  rather 
rounded  gametocytes.  According  to  such  an  observation  the  organism  would  belong 
to  Plasmodium  and  not  to  Laverania.  The  question  whether  the  tropical  fevers  are 
caused  by  two  different  parasites  does  not  seem  to  be  definitely  settled. 

The  young  trophozoite  of  the  malignant,  pernicious  tertian,  or  sub- 
tertian  parasite  (fig.  87)  are  but  slightly  active  and  are  very  small,  even 
after  the  formation  of  the  comparatively  large  food  vacuole,  which 
makes  the  body  appear  annular  (''signet  ring"  stage).  Often  two  and 
even  more  parasites  are  found  in  one  blood  corpuscle. 

Fully  grown  they  only  attain  two-thirds  or  less  of  the  diameter  of 
the  erythrocytes,  which  display  an  inclination  to  shrink  and  then  appear 


i68 


THE   ANIMAL   PARASITES   OF   MAN 


darker  than  the  normal  (brass-coloured).  In  the  early  stage  dots  or 
stippling — sometimes  called  Maurer's  dots — appear  on  the  blood  cor- 
puscles as  in  those  attacked  by  the  ordinary  tertian  parasite  {Plasmo- 
dium vivax),  but  the  Maurer's  dots  are  relatively  coarse  and  few,  and 
are  not  easily  stained.  These  dots  were  first  described  by  Stephens 
and  Christophers  in  1900,  and  subsequently  by  Maurer  in   1902. 

About  thirty  hours  after  the  entrance  into  the  blood  corpuscles, 
the  parasites  are  rarely  found  in  the  peripheral  blood,  but  they  are 
present  in  the  internal  organs,  and  especially  in  the  spleen.  The 
schizogony,  which  now  begins  in  the  internal  organs,  proceeds  on  the 
same  lines  as  that  of  the  quartan  parasite,  that  is,  usually  with  the 
merozoites  radially  arranged  around  a  central  agglomeration  of  dark 
brown  pigment. 


m 


Fig.  87. — The  pernicious  malignant  or  sub-tertian  parasite  in  the  red  corpuscles  of  man, 
asexual  stages.     (After  Manson.) 

The  number  of  merozoites  formed  is  quoted  differently,  e.g.,  8  to  24, 
on  an  average  12  to  16.  However,  according  to  the  recent  cultural 
researches  of  J.  G.  and  D.  Thomson^  (1913)  the  number  of  merozoites 
of  P.  falciparum  is  32.  D.  Thomson,  from  examination  of  spleen 
smears  at  autopsy,  also  concludes  that  the  number  of  merozoites  may 
reach  32.  During  their  formation  the  blood  corpuscle  which  is 
attacked  gets  paler  and  disintegrates. 


Fig.  88. — The  crescents  of  the  malignant  tertian  parasite. 

See  also  fig.  8i. 


(After  Mannaberg.) 


The  gametocytes  which  finally  appear  are  attenuated,  curved 
bodies,  rounded  at  each  end  and  known  as  crescents  (figs.  81,  88),  and 
are  provided  with  a  nucleus  and  with  coarse  pigment  masses.  In  the 
males  the  pigment  is  more  scattered  than  in  the  females,  where  it  is 
around  the  nucleus.     Their  length  is  9  yLt  to  14  /a,  and  their  breadth  is 


•  Proc.  Roy.  Soc,  B,  Ixxxvii,  p.  77. 


LAVERANIA   MALARIA  169 

2  yLt  to  3  /x.  At  first  they  are  still  in  the  pale  blood  corpuscles,  later 
they  free  themselves  and  are  found  in  numbers  in  the  peripheral 
blood  in  cases  of  pernicious  malaria  of  Southern  Europe  and  the 
tropics,  while,  on  the  other  hand,  they  occur  much  more  rarely  in 
the  peripheral  blood  in  West  African  malignant  tertian.  Their  further 
development  takes  place  under  the  same  conditions  as  in  the  other 
malarial  parasites. 


Fig.  89. — Section  through  a  tubule  of  the  salivary  gland  of  an  Anopheles  with  sporozoites 
of  the  malignant  tertian  parasites  ;  on  the  left  at  the  top  a  single  spore  zoite  greatly  magnified. 
(After  Grassi). 

D.  Thomson  (1914),^  from  studies  of  autopsy  smears,  has  shown 
that  crescents  develop  chiefly  in  the  bone-marrow  and  spleen,  and 
take  about  ten  days  to  grow  into  the  a'dult  state  in  the  internal  organs. 
He  believes  that  crescents  are  produced  from  ordinary  asexual  spores. 
Quinine,  he  states,  has  no  direct  destructive  action  on  crescents,  but 
it  destroys  the  asexual  source  of  supply. 

The  sporozoites  of  Laverania  malarice  (P.  falciparum)  are  repre- 
sented in  fig.  89. 

The  principal  distinctive  characters  of  the  malignant  tertian  para- 
site are  :  (i)  The  ring  forms  are  very  small,  occasionally  bacilliform, 
and  may  be  marginal  ("  accole  "  of  Laveran)  ;  (2)  the  larger  tropho- 
zoites are  often  ovoid,  and  about  one-third  or  one-half  of  the 
erythrocyte  in  size;  (3)  the  infected  red  cells  sometimes  show  coarse 

'  Antials  Trop.  Med,  and  farasitoL,  viii,  p.  85. 


170  THE   ANIMAL   PARASITES   OF   MAN 

stippling  (Maurer's  dots)  ;  (4)  the  gametocytes,  or  sexual  forms,  are 
crescentic  in   shape. 

].  W.' W.  Stephens  (1914)  has  described  a  new  malarial  parasite  of 
man  ;  it  is  called  Plasmodium  temie.  It  is  very  amoeboid,  with  scanty 
cytoplasm  and  much  chromatin,  sometimes  rod-like  or  irregular. 
The  parasite  was  described  from  a  blood-smear  of  an  Indian  child. 
The  creation  of  a  new  species  for  this  parasite  has  been  criticized  by 
Balfour  and  Wenyon,  and  by  Craig. 

Plasmodium  relictum,  Sergent,  1907. 
Syn.  :  Plasmodium  prcECOx,  Grass!  and    Feletti,    1890;   Plasmodium   danilewskyi, 
Gr.  et  Fel.,  1890  ;  HcBmamoeba  relicta,   Gr.  et  Fel.,  1891  ;  Proteosoma  grassii, 
Labbe,  1894. 

Hasmamoeboid,  pigment-producing,  malarial  parasites  are  often  found  in  birds. 
Like  the  human  malarial  parasites  they  have  been  variously  named.  Labbe  created 
the  genus  Proteosoma  for  them,  and  this  name  is  still  often  used  as  a  distinctive  one 
unofficially.  The  correct  name  is  stated  to  be  either  Plasmodium  relictum  or 
P.  prcscox^  or  possibly  even  P.  danilewskyi,  assuming  that  there  is  only  one  species. 
The  nomenclature  of  the  malarial  parasites  is  most  confused.  The  avian  malarial 
parasites  are  transmitted  by  Culicine  mosquitoes. 

The  organism  was  discovered  by  Grassi  in  the  blood  of  birds  in  Italy,  and  causes 
a  fatal  disease  in  partridges  in  Hungary.  Sparrows  are  affected  in  India,  and  it 
was  this  Plasmodium  in  which  Ross  first  traced  the  development  of  a  malarial 
parasite  in  a  mosquito.  The  parasite  may  be  transmitted  from  bird  to  bird  by 
blood-inoculation,  canaries  being  very  susceptible. 

The  principal  stages  of  the  avian  plasmodium  closely  resemble  those  of  the 
malarial  parasites  of  man.  In  its  earliest  stage  P.  relictum  is  unpigmented, 
but  soon  the  trophozoite  grows  and  becomes  pigmented,  meanwhile  displacing 
the  nucleus  of  the  avian  red-blood  corpuscle,  a  characteristic  feature,  distinguish- 
ing it  from  Halteridium.  Schizonts  are  formed,  each  of  which  gives  rise  to 
about  nine  merozoites  in  the  circulating  blood.  Sexual  forms  or  gametocytes  also 
occur  in  the  blood.  These  develop  in  Culex  fatigans,  C.  pipiens  and  C.  nemor- 
osus.  Ookinetes  or  vermicules  are  formed  in  twelve  to  fifteen  hours  in  the 
stomach  of  the  mosquito,  and  in  one  to  two  days  well-developed  round  oocysts  may 
be  seen.  In  three  to  four  days  sporoblasts  have  formed  within  the  oocysts  and 
young  sporozoites  begin  to  develop.  'In  nine  to  ten  days  the  oocysts  are  mature, 
being  filled  with  sporozoites.  The  oocysts  then  burst  and  the  sporozoites  travel 
through  the  thoracic  muscles  to  the  salivary  glands  of  the  Culicine. 

Neumann,  experimenting  with  canaries,  found  that  Stegomyia  fasciata  could 
transmit  the  infection,  but  less  efficiently  than  species  of  Culex. 

The  Cultivation  of  Malarial  Parasites. 
The  successful  cultivation  of  malarial  parasites  in  vitro  was  first 
recorded  by  C.  C.  Bass  and  by  Bass  and  Johns  (191 2) .^     Since  then, 
J.  G.and  D.  Thomson,^  and  McLellan  (1912-13),  Ziemann^  and  others 
have  repeated  the  experiments. 

'  Journ.  Exptl.  Med.,  xvi,  p.  567. 

*  Annals  Trop.  Med.  and  ParasitoL,  vi,  p.  449;  vii,  pp.  153,  509. 

^   Tratis.  Soc.  Trop.  Med.  and  Hyg.,  vi,  p.  220. 


MALARIAL   PARASITES  IJI 

DIFFERENTIAL   CHARACTERS    OF    THE   HUMAN    MALARIAL   PARASITES. 


Character 

Plastnodiutn  tnalatice 
(Quartan) 

Plas7nodiu77i  vj'vax 
(Benign  tertian) 

Laverania  tnalarice  = 

Plastnodiu  vt  falciparum 

(Malignant  tertian) 

Schizogony    ... 

Complete  in  seventy- 
two  hours 

Complete     in     forty- 
eight  hours 

Complete  in  forty-eight 
hours  or  less 

Trophozoite 

Smaller  than  P.vivax, 
larger       than      L, 
malaria 

Pseudopodia        n  o  t 
marked  or  long 

Young       trophozoite 
large. 

Long  pseudopodia 

Young  trophozoite 
small 

Movements 

Rather  slow  in  imma- 
ture forms 

Active      a  m  oe  b  o  i  d 
movements 

Sometimes  actively 
motile 

Pigment         

Coarse  granules,  peri- 
pherally   arranged, 
little  movement 

Fine    granules,    with 
active  movement 

Granules  fine  and 
scanty,  movement 
oscillatory 

Schizont         

Smaller  than  red  cor- 
puscle 

Larger       than       red 
blood   corpuscle 

Smaller  than  red 
corpuscle 

Merozoites     

6  to  12  forming  rosette 

15     to    20    regularly 
arranged 

8  to  32  (according  to 
different  authors)  ar- 
ranged  irregularly 

Gametocytes ... 

Spherical 

Spherical 

Crescentic 

Distribution   of  para- 
sites   in    vertebrate 
host 

About  equal  numbers 
in    peripheral    and 
visceral  blood 

Larger    numbers     in 
visceral  blood 

Scanty  in  peripheral 
blood  compared  with 
the  enormous  num- 
bers in  the  internal 
organs.  The  latter 
part  of  the  cycle 
(schizogony)  may 
occur  in  the  internal 
organs  only 

Alterations  in  erythro- 
cytes 

Almost  normal 

Pale     and    hypertro- 

phied. 
Schiiffner's  dots  seen 

in     deeply    stained 

specimens 

Corpuscle  may  be 
shrunken  and  dark, 
or  may  be  colourless. 
Maurer's  coarse  dots 
sometimes  seen 

Essentially  the  methoci  of  cultivation,  as  used  by  Thomson, 
is  as  follows  :  10  c.c.  of  infected  blood  are  drawn  from  a  vein  and 
transferred  to  a  sterile  test  tube,  in  which  is  a  thick  wire  leading  to 
the  bottom  of  the  tube.  One-tenth  of  a  cubic  centimetre  of  a 
50  per  cent,  aqueous  sokition  of  glucose  or  dextrose  is  placed 
in  the  test  tube,  preferably  before  adding  the  blood.  The  blood 
is  defibrinated  by  stirring  gently  with  the  wire.  When  defibrination 
is  complete  the  wire  and  the  clot  are  removed,  and  the  glucose-blood 
is  transferred,  in  portions,  to  several  smaller  sterile  tubes,  each  con- 
taining a  column  of  blood  about  one  inch  in  height.  The  tubes 
are  plugged  and  capped  and  then  transferred,  standing  upright,  to  an 


172  THE   ANIMAL   PARASITES   OF    MAN 

incubator  kept  at  a  temperature  of  37°  C.  to  41°  C.  The  blood  cor- 
puscles soon  settle,  leaving  a  column  of  serum  at  the  top,  to  the  extent 
of  about  half  an  inch  in  each  tube.  The  leucocytes  need  not  be 
removed  by  centrifugalization.  J.  G.  Thomson  (1913)  and  his 
collaborators  did  not  find  it  necessary  to  destroy  the  complement  in 
the  serum,  and  they  found  that  the  malarial  parasites  developed  at  all 
levels  in  the  column  of  corpuscles,  and  not  merely  on  the  surface 
layer  of  the  corpuscles  as  first  stated  by  Bass  and  Johns. 

So  far  only  the  asexual  generation  of  the  malarial  parasites  has 
been  grown  in  vitro.  Thomson  rarely  observed  haemolysis  in  the 
cultures.  Clumping  of  the  malignant  tertian  parasites  occurred. 
In  cultures  of  the  benign  tertian  parasite  {Plasmoditnn  z'^V^.r)  clumping 
was  not  observed.  J.  G.  and  D.  Thomson  consider  that  this 
difference  as  regards  clumping  explains  why  only  young  forms  of 
malignant  tertian  are  found  in  peripheral  blood,  as  the  clumping 
tendency  of  the  larger  forms  causes  them  to  be  arrested  in  the  finer 
capillaries  of  the  internal  organs.  It  also  explains  the  tendency  to 
pernicious  symptoms,  such  as  coma,  in  malignant  tertian  malaria. 
Further  it  was  found  from  cultures  that  P.  falciparum  was  capable 
of  producing  thirty-two  spores  (merozoites)  in  maximum  segmenta- 
tion, while  P.  vivax  produced  sixteen  spores  (merozoites)  as  a  rule, 
though  the  number  might  be  greater  than  sixteen.  (Quartan  parasites 
produce  eight  spores  or  merozoites  in  schizogony.) 

It  may  also  be  mentioned  here  that  Babesia  {Piroplasma)  canis  has 
been  successfully  cultivated  in  vitro  by  Bass's  method.  This  has  been 
accomplished  by  Thomson  and  Fantham,^  Ziemann,  and  Toyoda  in 
1913.  J.  G.  Thomson  and  Fantham  used  the  simplified  Bass  technique 
recorded  above,  namely,  infected  blood  and  glucose,  incubating  at 
37°  C.  In  one  of  the  B.  canis  cultures,  starting  with  heart  blood  of  a 
dog  containing  corpuscles  infected  with  one,  two,  or,  exceptionally, 
four  piroplasmata,  Thomson  and  Fantham  succeeded  in  obtaining  a 
maximum  of  thirty-two  merozoites  in  a  corpuscle.  The  cultures  are 
infective  to  dogs  and  sub-cultures  have  been  obtained. 


Family.     Piroplasmidae,   Franga. 

The  parasites  included  in  this  provisional  family  or  group  belong 
to  the  Haemosporidia.  They  are  minute  organisms,  sometimes 
amoeboid,  but  usually  possessing  a  definite  form.  They  are  endo- 
globular,  being  contained  within  mammalian  red  blood  corpuscles, 
but  they  produce  no  pigment.  The  true  Piroplasmata,  belonging 
to   the  genus  Babesia,  destroy  the  host  corpuscles,  setting  free  the 

'  Annals  Trap.  Med.  and  ParasitoL,  vii,  p.  621. 


PIROPLASMID^ 


^11 


haemoglobin,  which  is  excreted  by  the  kidneys  of  the  cow,  sheep, 
horse,  dog,  etc.,  acting  as  host  The  disease  produced,  variously  called 
piroplasmosis  or  babesiasis,  is  consequently  characterized  by  a  red 
coloration  of  the  urine  known  as  haemoglobinuria,  or  popularly  as 
^'red-water."  One  of  the  best  known  piroplasms  is  Piroplasma 
bigeiniiiiuji  or  Babesia  bovis  (probably  the  latter  name  is  correct), 
which  is  the  causal  agent  of  "  Texas  fever  "  or  '^  red-water  "  in  cattle 
and  is  spread  by  ticks. 


©showing  the  usual  mode 
of  multiplication 
of 
Nuttallia  equi 

in  the  circulating  blood 


Fig.  90. — Nuttallia  equi,  life-cycle  as  seen  in  red  blood  corpuscles  in  stained  preparations 
of  peripheral  blood.     (After  Nuttall  and  Strickland.) 


Of  recent  years,  researches  on  the  morphology  of  these  blood 
parasites  has  led  to  their  separation  into  various  genera  and  species. 
However,  our  knowledge  is  still  very  far  from  complete.     The  various 


174  THE   ANIMAL   PARASITES   OF   MAN 

genera  recognized  by  Fran9a^  (1909),  and  placed  in  a  provisional 
family,  Piroplasmidai,  may  be  listed,  though  further  research  may 
lead  to  emendations  : — 

(i)  Babesia  (Starcovici)  or  Piroplasma  (Patton).  Pyriform 
parasites,  dividing  by  a  special  form  of  budding  or  gemmation  with 
chromatin  forking,  as  well  as  by  direct  binary  fission.  Parasitic  in 
oxen,  dogs,  sheep,  horses,  etc. 

(2)  rheileria  (Bettencourt,  Fran9a  and  Borges).  Rod-shaped  and 
oval  parasites  occurring  in  cattle  and  deer.  T.  parva  is  the  patho- 
genic agent  of  African  East  Coast  fever  in  cattle. 

(3)  NiittaUla  (Fran9a).  Oval  or  pear-shaped  parasites,  with  mul- 
tiplication in  the  form  of  a  cross.  iV.  equi''  (fig.  90)  of  equine  ''piro- 
plasmosis"  (nuttalliosis).     A^.  herpestidis  in  a  mongoose. 

(4)  NicoUia  (Nuttall).  Oval  or  pear-shaped  parasites  with 
characteristic  nuclear  dimorphism,  and  with  quadruple  division  at 
first  fan-like,  then  like  a  four-leaved  clover.  A^.  qiiadrigemina  from 
the  gondi. 

(5)  Smithia  (Fran9a).  Pear-shaped,  single  forms  stretching  across 
the  blood  corpuscle.  Multiplication  into  four  in  the  form  of  a  cross. 
S.  microti  from  Microtus  arvalis,  S.  talpce  from  the  mole. 

(6)  Rossiella  (Nuttall).  This  belongs  to  the  family  Piroplasmidse 
of  Fran9a.  It  is  intracorpuscular  and  non-pigment  forming,  occurring 
singly,  in  pairs,  or  occasionally  in  fours.  It  is  usually  round  and 
larger  than  Babesia.  The  parasite  multiplies  by  binary  fission. 
R.  rossi  in  the  jackal. 

The  genus  Babesia  is  the  best  known  and  most  important,  and 
will  be  considered  next. 

Genus.     Babesia,  Starcovici,   1893. 

Syn.  :  Pyrosoma,  Smith  and  Kilborne,  1893  >  Apiosojna^  Wandolleck,   1895  '  Piro- 
plasma, W.  H.  Patton,  1895  ;  AmcEbosporidium^  Bonome,  1895. 

The  organisms  belonging  to  this  genus  are  pyriform,  round  or 
amoeboid.  The  characteristic  mode  of  division  is  as  follows :  Just 
before  division  the  parasite  becomes  amoeboid  and  irregular  in  shape, 
(fig.  91,  I — 5)  with  a  compact  nucleus.  The  latter  gives  off  a  nuclear 
bud.  This  nuclear  bud  divides  into  two  by  forking  (fig.  91,  6,  7) 
The  chromatin  forks  grow  towards  the  surface  of  the  body  of  the 
rounded  parasite,  and  then  two  cytoplasmic  buds  grow  out.  The 
forking  nuclear  buds,  which  are  Y-shaped,  pass  into  the  cytoplasmic 
outgrowths^  (fig.  91,  S,   9).     The  buds  gradually  increase  in  size  at 

•  Arch.  Inst.  Bact.  Camara  Pestatta,  iii,  p.  ii. 

'^  Parasitology,  v  (IQ12),  p.  65. 

^  Nuttall  and  Graham-Smith,  Journ.  Hyg.,  vii,  p.  232. 


BABESIA 


175 


showing  the  usual  mode 
of  multiplication 

of 

Piroplasma  canis 

in  the  circulating  blood. 


15 


Fig.  91. — Babesia  {Piroplasma)  canis,  life-cycle  in  stained  preparations  of  infected  blood 
of  dog.     (After  Nuttall  and  Graham-Smith.) 


176  THE   ANIMAL   PARASITES   OF   MAN 

the  expense  of  the  parent  form  until  they  become  two  pear-shaped 
parasites  joined  at  their  pointed  ends.  The  connecting  strand  shrinks 
and  the  two  daughter  forms  separate  (fig.  91,  10 — 14).  The  pyriform 
parasites  after  having  exhausted  the  blood  corpuscle  escape  from  it 
(fig.  91, 15),  and  seek  out  fresh  host  corpuscles,  entering  by  the  rounded, 
blunt  end  (fig.  91,  i).  It  is  the  pyriform  phase  of  the  parasite  which 
penetrates  red  blood  corpuscles,  not  rounded  forms,  which  die  if  set 
free.  The  pyriform  parasite,  however,  becomes  rounded  (fig.  91,  2,  3), 
soon  after  its  entry  into  a  fresh  host  cell.  This  interesting  mode  of 
division  by  gemmation  and  chromatin  forking  has  been  made  dia- 
gnostic of  the  genus  Babesia  by  Nuttall.^  Rounded  forms  of  Babesia 
divide  by  binary  fission,  and  this  direct  method  can  also  be  adopted 
by  the  other  forms  of  Babesia. 

The  distribution  of  the  chromatin  in  the  pear-shaped  Babesia,  as  seen  in  B.  cams 
and  B.  bovis,  is  interesting.  The  main  nuclear  body  consists  of  a  karyosome 
surrounded  by  a  clear  area.  There  is  also  a  loose  (chromidial)  mass  of  chromatin 
representing  the  remains  of  the  chromatin  forks  seen  during  the  formation  of  the 
parasite  as  a  daughter  form  by  gemmation.  Occasionally  there  is  a  small  dot  or 
point,  the  so-called  "  blepharoplast"  of  Schaudinn  and  Liihe.  This  minute  dot  is 
not  a  flagellate  blepharoplast,  for  there  is  no  flagellate  stage  in  the  life-history  of 
Babesia.  These  nuclear  phenomena  have  been  described  by  Nuttall  and  Graham- 
Smith  and  Christophers  (1907)^  for  B.  cants,  by  Fantham  (1907)^  for  B.  bovis,  and 
by  Thomson  and  Fantham  (1913)  from  glucose-blood  cultures  of  ^.  cams. 

Babesia  are  tick  borne,  as  was  first  shown  by  Smith  and  Kilborne 
(1893).  The  developmental  cycle  in  the  tickis  in  completely  known. 
The  best  accounts  are  those  of  Christophers  (1907)'*  for  B.  cams  and 
Koch  (1906)  for  B.  bovis,  and  these  accounts  are  supplementary. 
The  principal  stages,  so  far  as  known,  may  be  summarized  thus  : — 

(i)  The  piroplasms  taken  by  the  tick  in  feeding  on  blood  pass  into  the  tick's 
stomach.  The  pyriform  parasites,  which  alone  are  capable  of  further  development, 
are  set  free  from  the  blood  corpuscles.  In  about  twelve  to  eighteen  hours  they 
become  amoeboid,  sending  out  long,  stiff,  slender,  pointed  pseudopodia.  The 
nucleus  of  each  parasite  divides  unequally  into  two.  Similar  forms  have  been 
obtained  in  cultures.  These  stellate  forms  may  be  gametes,  and  according  to  Koch 
fuse  in  pairs. 

(2)  A  spherical  stage  follows,  possibly  representing  the  zygote.  This  grows,  and 
a  uninucleate  globular  mass  results.  This  form  is  found  in  large  numbers  on  the 
third  day,  according  to  the  observations  of  Koch. 

(3)  A  club-shaped  organism  is  next  formed.  This  may  represent  an  ookinete 
stage.  The  club-shaped  bodies  are  motile  and  gregarine-like,  and  are  about  four 
times  the  size  of  the  blood  forms.  These  club-shaped  bodies  and  subsequent  stages 
were  described  by  Christophers  in  the  development  of  B.  cams  in  the  dog-tick, 
Rhipicephalus  sanguineus. 

'  "  Piroplasmosis,"  Herter  Lectures,  Parasitology,  vi,  p,  302. 

"^  Sci.  Mems.  Govt.  India,  No.  29. 

3  Quart.  Journ.  Microsc.  Sci.,  li,  p.  297. 

*  Sci.  Mems.  Govt.  India,  No.  29. 


BABESIA  177 

(4)  The  club-shaped  bodies  pass  from  the  gut  of  the  tick  into  the  ovary,  and  so 
get  into  the  ova.  There  they  become  globular,  and  later  are  found  in  the  cells  of 
the  developing  tick-embryo.  The  parasites  are,  then,  transmitted  hereditarily. 
Similar  globular  bodies  are  found  in  the  tissue  cells  of  the  body  of  tick  nymphs 
which  have  taken  up  piroplasms.  The  globular  stage  was  called  the  "zygote"  by 
Christophers,  but  it  may  correspond  to  the  oocyst  of  Plasmodia. 

(5)  The  globular  body  divides  into  a  number  of  "sporoblasts,"  which  become 
scattered  through  the  tissues  of  the  larval  or  nymphal  tick,  as  the  case  may  be. 

(6)  The  sporoblasts  themselves  divide  into  a  large  number  of  sporozoites,  which 
are  small  uninucleate  bodies,  somewhat  resembling  blood  piroplasms.  The  sporo- 
zoites collect  in  the  salivary  glands  of  the  tick.  They  are  inoculated  into  the 
vertebrate  when  the  tick  next  feeds. 

The  chief  species  of  Babesia  and  their  pathogenic  importance 
may  be  listed  thus  : — 

(i)  Babesia  bovis  (Babes)  produces  infectious  haemoglobinuria 
of  cattle  in  Europe  and  North  Africa.  It  is  transmitted  by  Ixodes 
riciiiits.     A  similar  parasite  also  occurs  in  deer. 

(2)  Babesia  bigeniina  (Smith  and  Kilborne)  produces  Texas 
fever,  tristeza,  or  red-water  in  cattle  in  North  and  South  America, 
South  Africa  and  Australia.  It  is  transmitted  by  Bodphilns  annulatiis 
in  North  America,  by  B.  austral  is  in  Australia,  South  America,  and 
the  Philippines,  and  by  B.  decoloratus  in  South  Africa. 

The  parasite  is  from  2  /a  to  4  /x  long,  and  from  i'5  yu,  to  2  yu,  broad. 
Babesia  bigemina  may  be  the  same  parasite  as  B.  bovis. 

(3)  Babesia  divergens  (MacFadyean  and  Stockman)  is  a  small 
parasite.  It  is  found  in  cattle  suffering  from  red-water  in  Norway, 
Germany,  Russia,  Hungary,  Ireland,  Finland,  and  France,  and  is 
transmitted  by  Ixodes  ricinus. 

(4)  Babesia  canis  (Plana  and  Galli-Valerio)  gives  rise  to  malignant 
jaundice  or  infectious  icterus  in  dogs  in  Southern  Europe,  India,  and 
other  parts  of  Asia  and  North  Africa,  where  it  is  transmitted  by 
Rhipicephalus  sanguineus.  In  Africa  generally,  especially  South 
Africa,  the  disease  is  transmitted  by  Hceinaphysalis  leacJii.  Babesia 
canis  varies  from  07  /a  to  5  //-,  the  size  depending  partly  on  the 
number  of  parasites  w^ithin  the  corpuscle.  It  averages  about  3  //,.  It 
has  been  cultivated  in  Bass'  medium  (glucose  and  infected  blood), 
see  p.  172. 

In  India  Piroplasma  gibsoni  (Patton)  infects  hunt  dogs  and  jackals. 
It  is  annular  or  oval  in  shape. 

(5)  Babesia  ovis  (Babes)  produces  ^'  Carceag,"  a  disease  of  sheep 
in  Roumania,  the  Balkan  Peninsula,  Italy,  and  Transcaucasia.  It 
varies  in  size  from  i  />«,  to  3  //,.  It  is  transmitted  by  Rhipicephalus 
bursa.     The  parasite  has  recently  been  recorded  from  Rhodesia. 

(6)  Babesia  caballi  (Nuttall  and  Strickland)  causes  *'  biliary 
fever"  in   equines.     The  parasite   occurs  in    Russia,   Roumania,   and 

12 


iy8  THE   PARASITES   OF   MAN 

Transcaucasia.     It  varies  in  size  from  j  /jl  to  2  fi.     It  is  transmitted  by 
Dermacentor  reticulaius. 

It  should  be  mentioned  that  Nuttallia  equi  also  causes  "piroplasmosis"  in 
equines,  with  symptoms  of  hasmo^lobinuria  and  jaundice  in  Italy,  Sardinia,  many 
parts  of  Africa,  Tianscaucasia,  India,  and  Brazil.  In  Africa  it  is  transmitted  by 
Rhipicephalus  evertsi.  It  has  been  shown  experimentally  that  a  horse  recovered 
from  Babesia  caballi  was  susceptible  to  the  inoculatit)n  of  Nuttallia  equi  blood. 

(7)  Babesia  pitheci  (P.  H.  Ross)  was  found  in  a  monkey, 
Cercopithecus  sp.,  in  Uganda.  The  pear-shaped  forms  measure  1*5  fi 
by  2*5  fjb. 

(8)  Babesia  inuris  (Fantham)^  was  found  in  w^hite  rats.  The 
pyriform  parasites  are  2  fi  to  ^  /jl  long  and  i  ft  to  i'5  fju  broad;  oval 
forms  are  0-5  to  1*5  fi  diameter. 

The  usual  symptoms  of  babesiasis  (piroplasmosis)  are  high  fever, 
loss  of  appetite,  haemoglobin iiria,  icterus,  anaemia,  paralysis,  and 
death  in  about  a  week  in  acute  cases.  In  chronic  cases  there  is 
anaemia,  and  haemoglobinuria  is  less  marked.  When  animals  recover, 
there  are  still  some  piroplasms  left  in  the  blood.  "  Recovered  "  or 
*' salted"  animals  are  not  susceptible  to  reinfection,  but  ticks  feeding 
on  them  acquire  piroplasms,  and  are  a  source  of  danger  to  freshly 
imported  animals. 

Treatment. — Trypan-blue  is  the  best  drug,  as  shown  by  Nut  tall  and  Had  wen* 
(1909).  It  should  be  administered  intravenously  in  i  to  1*5  per  cent,  aqueous  solu- 
tion. A  dose  of  5  to  10  c.c.  is  curative  for  dogs,  one  of  100  to  150  c.c.  for  horses  and 
cattle.  Unfortunately,  the  tissues  are  coloured  blue  by  the  drug.  The  "salted" 
animals,  after  trypan-blue  treatment,  still  harbour  the  parasites  in  their  blood  for 
years. 

Genus.     Theileria,  Bettencourt,  Franqa  and  Borges,  1907. 

The  organisms  belonging  to  this  genus  are  rod-like  or  bacilliform,  and  coccoid 
or  round. 

The  best  known  of  the  species  of  Theileria  is  T.  parva^  the  pathogenic  agent  of 
East  Coast  fever  or  Rhodesian  fever  in  cattle  in  Africa. 

Theileria  parva,  Theiler,  1903. 

Syn.  :  Piroplasma  farvum. 
In  the  blood  corpuscles  of  infected  cattle  minute  rod-like  and 
oval  parasites  are  seen.  Some  are  comma  siiaped  and  otht-rsare 
clubbed  (fig.  92,  1-12).  The  rod-like  forms  measure  i  /^  to  3  /^  in 
length  by  0*5  yu<  in  breadth;  the  oval  forms  are  0*7  //,  to  1-5  /a  in 
diameter.  The  intracorpuscular  parasites  are  said  by  R.  Gonder 
{1910)  to  be  gametocytes,  the  rod-like  forms  being  thought  to 
be  males,  the  oval  forms  to  be  females.  Free  parasites  are  practi- 
cally never  seen  in  the  blood.     It  is  known  that  it  is  impossible  to 

'  Quart,  fourn.  Microsc.  Sci.,  1,  p.  493. 
2  Parasitology,  ii,  p.  156. 


THEILERIA    PARVA 


179 


produce  the  disease  in  a  healthy  animal  by  blood  inoculation,  but 
only  by  intraperitoneal  transplantation  of  large  pieces  of  infected 
spleen  (Meyer).  There  may  be  as  many  as  eight  parasites  in  a  cor- 
puscle. The  chromatin  is  usually  at  one  end  of  the  organism.  In 
some  parasites  the  appearance  of  the  chromatin  suggests  division,  but 
such  division,  if  it  takes  place,  must  be  very  slow,  as  it  has  not  been 
actually  seen  in  progress.  The  red  blood  corpuscles  appear  merely  to 
act  as  vehicles  for  the  parasites  (Nuttall,  Fantham,  and  Porter).^ 


00© 


14 


16 


Fig.  92. —  Theileria  parva.  1-12,  intracorpuscular  parasites,  stained.  (After  Nuttall  and 
Famham)  ;  13-18,  Koch's  blue  bodies,  from  stained  spleen  smear;  17-18,  breaking  up  of 
Koch's  body.     (After  Nuttall.) 

In  the  internal  organs,  especially  the  lymphatic  glands,  spleen  and  bone-marrow, 
are  found  multinucleate  bodies  known  as  Koch's  blue  bodies  (fig.  92,  13-18).  These 
are  schizonts,  according  to  Gonder.^  The  actual  Koch's  blue  bodies  are  said  to  be 
extracellular,  but  similar  multinucleate  bodies,  schizonts,  occur  in  lymphocytes.  The 
schizonts  divide  and  the  merozoites  resulting  probably  invade  the  red  blood  cor- 
puscles in  the  internal  organs.  Gonder  considers  that  the  sporozoites  injected  by 
the  tick  collect  in  the  spleen  and  lymphatic  glands,  penetrate  the  lymphocytes  and 
give  rise  to  the  schizonts. 

Gonder  has  studied  the  cycle  of  7 .  parva  in  the  tick.  He  states  that  the 
gametocytes  leave  the  host  corpuscles  and  give  rise  to  gametes,  then  conjugation 
occurs  producing  zygotes.  The  zygotes  are  then  said  to  become  active  lo  form 
ookinetes,  and  to  enter  the  salivary  glands  of  the  tick.  Multiplication  is  said  to 
occur  therein,  producing  a  swarm  of  sporozoites.     This  work  needs  confirmation. 

T.  parva  is  transmitted  by  Rhipicephalus  appendiculatus^  R.  simus,  R.  evertsi^ 
R.  m/ens,  and  R.  capensis.  The  parasites  are  not  hereditarily  transmitted  in 
Rhipicephalus^  but  when  taken  by  the  transmitter  at  one  stage  of  its  development 
the  tick  is  infective  in  its  next  stage  (<?.^.,  if  the  larva  becomes  infected,  then  the 
nymph  is  infective ;  if  the  nymph  becomes  infected,  then  the  adult  is  infective). 

An  animal  recovered  from  Theileria  parva  is  incapable  of  infecting  ticks,  but  few 
animals  recover  from  East  Coast  fever.  Animals  suffering  therefrom  do  not  show 
haemoglobin  uria. 


'  Farasitology^  ii,  p.  325  ;  iii,  p.  117. 

'^  Zeitschr.f.  Infekt.  paras.  Krankh.  u.  Hyg.  d.  Hausttere,  viii,  p.  406. 


l8o  THE   ANIMAL   PARASITES   OF   MAN 

Theileria  mutans,  Theiler^   'Qoy. 
Syn.  :  Piroplasma  miitans. 
This  is  transmissible  experimentally  by  blood  inoculation.     It  occurs  in  cattle  in 
South  Africa  and  Madagascar  and  is  apparently  non-pathogenic.     No  Koch's  blue 
bodies  are  formed.     It  is  transmitted  by  ticks. 

Theileria  amiulata  (Dschunkowsky  and  Luhs)  occurs  in  cattle  in  Transcaucasia. 
A  Theileria  {T.  stordii)  has  been  found  in  a  gazelle  (Franqa,  191 2). 

Genus.  Anaplasma,  Theiler,  1910. 
This  genus^  may  be  mentioned  here.  The  organisms  included  therein  are, 
according  to  Theiler,  coccus-like,  consisting  of  chromatin,  and  are  devoid  of  cyto- 
plasm. They  occur  in  the  red  blood  corpuscles  of  cattle,  causing  a  disease  charac- 
terized by  destruction  of  red  cells,  fever  and  anaemia,  but  with  yellow  urine.  The 
disease  is  tick  transmitted.  The  bodies  now  called  Anaplas7na  mar^inale  were 
formerly  described  as  marginal  points.  They  multiply  by  simple  fission.  They  are 
said  by  Theiler  to  cause  gall-sickness  in  cattle  in  South  Africa.  Some  authors  doubt 
whether  these  bodies  are  organismal. 

Genus.     Paraplasma,  Seidelin,  191 1. 

Under  this  generic  name  Seidelin  described  certain  bodies  found 
by  him  in  cases  of*  yellow  fever  in  1909.  The  type  species  is 
P.  flavigetiiim,^  and  is  claimed  by  Seidelin  to  be  the  causal  agent  of 
yellow  fever. 

Paraplasma  flavigemiin  occurs  in  the  early  days  of  the  disease  as 
small  chromatin  granules  with  or  without  a  faint  trace  of  cytoplasm. 
The  bodies  are  usually  intracorpuscular.  Also,  somewhat  larger  forms, 
with  distinct  cytoplasm,  are  seen  in  small  numbers.  During  the  later 
days  of  the  disease  still  larger  forms  are  found,  and  these  occur  also 
in  sections  of  organs  {e.g.,  kidney)  made  post-mortem.  Some  of  these 
larger  forms  are  perhaps  schizonts.  In  the  second  period  of  the 
disease  possible  micro-  and  macro-gametes  may  be  found,  some  of 
which  are  extracorpuscular.  Some  small  free  bodies  have  been  seen. 
Recently  schizogony  has  been  stated  to  occur  in  the  lungs,  and  it  is 
said  that  guinea-pigs  can  be  inoculated  with  Paraplasma  flavigeimm, 
and  show  yellow  pigment  in  the  spleen. 

Seidelin  places  Paraplasma  in  the  Babesiida^,  wath  resemblances 
more  particularly  to  Theileria.  V.  Schilling-Torgau  and  Agramonte 
have  criticized  these  findings  ;  the  former  considers  them  to  be  the 
resultant  of  certain  blood  conditions. 

P.  siihflavigemun  was  found  by  Seidelin  in  191 2  in  a  man  suffering 
from  an  unclassified  fever  in  Mexico. 

Further,  it  is  now  known  that  a  Paraplasma  occurs  naturally  in 
guinea-pigs.  More  researches  are  needed  on  these  matters,  as  some 
writers  {e.g.,  Wenyon  and  Low)  claim  that  the  bodies  are  not 
organismal. 

'  Bull.  Soc.  Path.  Exot.,  ill,  p.  135.  "^  Yellow  Fever  Bulletin,  i,  p.  251. 


NEOSPORIDIA  i8l 


Sub-class.     NEOSPORIDIA,  Schaudinn. 

Sporozoa   in    which  growth  and   spore   formation   usually   go    on 
together. 

Order.     Myxosporidia,  Butschli. 

These  parasites,  which  were  discovered  by  Johannes  Miiller  (1841),  Hve  principally 
in  fishes,  and  occasionally  cause  destructive  epizootics  amongst  their  hosts.  Miiller 
first  observed  them  in  the  form  of  whitish-yellow  pustules  on  the  skin  or  on  the  gills 
of  various  fishes.  These  pustules  contained  masses  of  small  shell-covered  bodies  with 
or  without  tails  ("psorosperms,"  see  fig.  93).  Similar  bodies  were  also  found  in  the 
air  bladders  of  certain  fish.  Creplin  (1842)  demonstrated  the  resemblance  of  the  cysts 
("psorosperm  tubes")  harbouring  the  psorosperms  to  the  "  pseudonavicella-cysts  "  of 
a  gregarine,  as  described  by  v.  Siebold.  Dujardin  (1845)  considered  that  there  was 
possibly  some  connection  between  the  protoplasmic  "psorosperm  tubes"  and  the 
spores  they  contained,  and  the  developmental  stages  of  monocystid  gregarines  from  the 


Paraplasma  flavigeiutni. — The  Yellow  Fever  Commission  (West 
Africa)  in  their  third  report,  dated  1915,  have  come  to  the  conclusion 
that  there  is  no  evidence  that  the  bodies  termed  Paraplasma  flavi- 
geiiitni  are  of  protozoal  nature  or  that  they  are  the  causal  agents  of 
yellow  fever. 


Face  p.   180.  

(natural  size),  with  two  myxospo-  7Wyxo^omJWmnerf,mm tne poiar 

ridia.    Lower  figures,  a,  d,  d,  spores  bodies  and  their  nuclei  and  the 

of  myxosporidia  from  a  pike,  Esox  sporozoite.     (After  Butschli.) 
lucius.    c,  Spore   from   Plaiy stoma 
fasciatiun.     (After  J.  Miiller.) 

placed  on  a  firmer  basis  by  Leydig  (185 1)  and  Lieberkiihn.  The  former  found  numerous 
forms  in  marine  fish,  and  he  discovered  in  species  which  live  free  in  the  gall  bladder 
of  cartilaginous  fishes  that  the  psorosperms  originated  in  a  manner  similar  to  the 
gregarines.  Lieberkiihn  (1854)  studied  the  Myxosporidia  in  the  bladder  of  the  pike 
(fig.  93,  a,  b,  cV),  and  observed  their  amoeboid  movements,  as  well  as  the  formation  of 
the  spores,  from  each  of  which  a  small  amoeboid  body  escaped,  a  discovery  that 
was  confirmed  by  Balbiani.  The  same  author  also  found  that  spiral  filaments  were 
enclosed  in  the  so-called  polar  body,  i.e.,  the  polar  capsule  of  the  psorosperm  spores, 
and  that  these  could  be  protruded  (fig.  93,  d,  and  fig.  95). 

The  term  Myxosporidia,  which  at  the  present  day  is  universally  applied  to  the 
"psorosperm  tubes,"  was  introduced  by  Butschli  in  1881,  who  studied  not  only  the 
structure  and   development  of  the   spores,  but  also  the  protoplasmic  body  of  the 


l8o  THE    ANIMAL   PARASITES   OF   MAN 

Thelleria  mutans,  Theiler,   IQO?- 
Syn.  :  Piroplasma  mutans. 
This  is  transmissible  experimentally  by  blood  inoculation.     It  occurs  in  cattle  in 
South  Africa  and  Madagascar  and  is  apparently  non-pathogenic.     No  Koch's  blue 
bodies  are  formed.     It  is  transmitted  by  ticks. 

Theileria  annulata  (Dschunkowsky  and  Luhs)  occurs  in  cattle  in  Transcaucasia. 
A  Theileria  {T.  stordii)  has  been  found  in  a  gazelle  (Franca,  1912). 

Genus.     Anaplasma,  Theiler,  1910. 

This  genus'  may  be  mentioned  here.  The  organisms  included  therein  are, 
according  to  Theiler,  coccus-like,  consisting  of  chromatin,  and  are  devoid  of  cyto- 
plasm. They  occur  in  the  red  blood  corpuscles  of  cattle,  causing  a  disease  charac- 
terized by  destruction  of  red  cells,  fever  and  anaemia,  but  with  yellow  urine.  The 
disease  is  tick  transmitted.  The  bodies  now  called  Anaplasma  mar^inale  were 
formerly  described  as  marginal  points.  They  muUiply  by  simple  fission.  They  are 
said  by  Theiler  to  cause  gall-sickness  in  cattle  in  South  Africa.  Some  authors  doubt 
whether  these  bodies  are  organismal. 


Recently  schizogony  has  been  stated  to  occur  in  the  lungs,  and  it  is 
said  that  guinea-pigs  can  be  inoculated  with  Paraplasma  flavigenuiu, 
and  show  yellow  pigment  in  the  spleen. 

Seidelin  places  Paraplasma  in  the  Babesiida',  with  resemblances 
more  particularly  to  Theileria.  V.  Schilling-Torgau  and  Agramonte 
have  criticized  these  findings  ;  the  former  considers  them  to  be  the 
resultant  of  certain  blood  conditions. 

P.  subflavigennin  was  found  by  Seidelin  in  191 2  in  a  man  suffering 
from  an  unclassified  fever  in  Mexico. 

Further,  it  is  now  known  that  a  Paraplasma  occurs  naturally  in 
guinea-pigs.  More  researches  are  needed  on  these  matters,  as  some 
writers  {e.g.,  Wenyon  and  Low)  claim  that  the  bodies  are  not 
organismal. 

'  Bji//.  Soc.  Path.  Exof.,  ill,  p.  135.  '^  Yellow  Fever  Bulletin,  i,  p.  251. 


NEOSPORIDIA 


l8l 


Sub-class.     NEOSPORIDIA,  Schaudinn. 


Sporozoa  in 
together. 


which  growth  and 


spore  formation   usually   go    on 


Order.     Myxosporidia,  Biitschli. 


These  parasites,  which  were  discovered  by  Johannes  Miiller  (1841),  live  principally 
in  fishes,  and  occasionally  cause  destructive  epizootics  amongst  their  hosts.  Muller 
first  observed  them  in  the  form  of  whitish-yellow  pustules  on  the  skin  or  on  the  gills 
of  various  fishes.  These  pustules  contained  masses  of  small  shell-covered  bodies  with 
or  without  tails  ("psorosperms,"  see  fig.  93).  Similar  bodies  were  also  found  in  the 
air  bladders  of  certain  fish.  Creplin  (1842)  demonstrated  the  resemblance  of  the  cysts 
("psorosperm  tubes")  harbouring  the  psorosperms  to  the  " pseudonavicella-cysts  "  of 
a  gregarine,  as  described  by  v.  Siebold.  Dujardin  (1845)  considered  that  there  was 
possibly  some  connection  between  the  protoplasmic  "psorosperm  tubes"  and  the 
spores  they  contained,  and  the  developmental  stages  of  monocystid  gregarines  from  the 
vesiculae  seminales  of  earth-worms.     The  relationship  of  the  "  fish  psorosperms  "  was 


a  b      c  d 

Fig.  93. — Upper  figure,  part  of 
a  gill  of  a  roach,  Leuciscus  rntihis 
(natural  size),  with  two  myxospo- 
ridia. Lower  figures,  a,  b,  d,  spores 
of  myxosporidia  from  a  pike,  Esox 
lucius.  c,  Spore  from  Platystoma 
fasciattim.     (After  J.  Miiller.) 


Fig.  94. — The  tailless  spore  of 
Myxobolus  miilhriyW\\\i\}[\Q  polar 
bodies  and  their  nuclei  and  the 
sporozoite.     (After  Biitschli.) 


placed  on  a  firmer  basis  by  Leydig  (185 1)  and  Lieberkiihn.  The  former  found  numerous 
forms  in  marine  fish,  and  he  discovered  in  species  which  live  free  in  the  gall  bladder 
of  cartilaginous  fishes  that  the  psorosperms  originated  in  a  manner  similar  to  the 
gregarines.  Lieberkiihn  (1854)  studied  the  Myxosporidia  in  the  bladder  of  the  pike 
(fig.  93,  a,  b,  cl)^  and  observed  their  amoeboid  movements,  as  well  as  the  formation  of 
the  spores,  from  each  of  which  a  small  amoeboid  body  escaped,  a  discovery  that 
was  confirmed  by  Balbiani.  The  same  author  also  found  that  spiral  filaments  were 
enclosed  in  the  so-called  polar  body,  i.e.^  the  polar  capsule  of  the  psorosperm  spores, 
and  that  these  could  be  protruded  (fig.  93,  d,  and  fig.  95). 

The  term  Myxosporidia,  which  at  the  present  day  is  universally  applied  to  the 
"psorosperm  tubes,"  was  introduced  by  Biitschli  in  1881,  who  studied  not  only  the 
structure  and   development  of  the   spores,  but  also  the  protoplasmic  body  of  the 


l82 


THE   ANIMAL   PARASITES   OF   MAN 


parasites  (fig.  96),  and  confirmed  the  occurrence  of  numerous  nuclei.  Many  authors 
have  made  important  additions  to  our  knowledge  of  the  Myxosporidia  :  Perugia, 
Thelohan,  Mingazzini,  L.  Pfeiffer,  L.  Cohn,  Doflein,  Mercier,  Schroder  and  Auerbach  ; 
while  the  presence  of  this  parasite  outside  the  class  of  fishes  has  become  known 
through  Lutz,  Laveran,  and  others.  The  species  causing  disease  in  fishes  have  been 
described  by  Ludwig,  Railliet,  Weltner,  L.  Pfeiffer,  Zschokke,  Hofer,  Doflein,  Gurley, 
Plehn,  Schuberg,  Fantham  and  Porter.  With  regard  to  classification  the  works  of 
Thelohan  (1895)  and  Gurley  (1894)  may  be  mentioned. 

The  Myxosporidia  live  either  free  on  the  epithelial  surface  of  hollow  organs 
(gall  or  urinary  bladder,  renal  tubules,  but  never  in  the  intestine),  or  are  enclosed  in 
the  tissues  of  their  host.  The  gills  and  muscular  system  are  their  favourite 
habitat,  but  other  tissues  or  organs  may  be  attacked.  Species  of  Myxosporidia 
are  also  known  from  Amphibia,  Reptilia,  and  a  few  invertebrates. 

The  free  forms,  which  are  often  amoeboid  (fig.  96),  move  by  the  aid  of  variously 
shaped  pseudopodia,  have  a  constant  form,  or  may  exhibit  contractions  of  the  body. 
The  tissue  parasites  often  reach  a  considerable  size,  so  that  the  integument  of  the 


p.s. 


Fig.  95. — Schematic  representation  of  a 
spore  of  Myxobolus.  One  polar  capsule  has 
protruded  its  filament ;  two  nuclei  and  a 
*'  vacuole  "  in  the  sporozoite.    (After  Doflein.) 


end 


Fig.  96. —  Chloromyxwn  leydigi.  Active 
trophozoite  (parasitic  in  gall-bladder  of  skates, 
rays,  dog-fish).  Ed,  ectoplasm  ;  ps^  pseudo- 
podia ;  end.  endoplasm  ;  y,  yellow  globules 
in  endoplasm ;  sp,  spores,  each  with  four 
polar  capsules,      x  525.     (After  Thelohan.) 


host  forms  protuberances  over  them.     They  are  of  a  roundish  or  irregular  shape 
Frequently  they  are  enveloped  in  a  connective  tissue  covering  formed  by  the  host. 

The  protoplasmic  body  in  the  trophic  phase  (fig.  96)  shows  a  distinct  ectoplasm 
which  is  finely  granular  or  sometimes  striated,  and  an  endoplasm  which  is  coarsely 
granular  and  contains  many  nuclei  as  well  as  cell  inclusions,  such  as  crystals,  pig- 
ment grains  and  fat  globules.  The  nuclei  originate  by  division  from  the  primitive 
nucleus  of  the  amoeboid  germ  that  issues  from  the  spore.  This  amoebula  may  or 
may  not  live  intra-cellularly  during  the  early  stages  of  its  existence. 

The  multinucleate  trophozoite  of  a  Myxosporidian  forms  spores  in  its  endoplasm 
practically  throughout  its  whole  period  of  growth  (fig.  96).  Vegetative  reproduction 
by  a  process  of  external  budding  or  plasmotomy  may  also  occur,  as  in  Myxidium 
lieberkiikm  from  the  urinary  bladder  of  the  pike. 

The  myxosporidian  trophozoite  may  produce  two  spores  within  itself,  when 
it  is  placed  in  the  sub-order  Disporea^  or  it  may  produce  numerous  spores,  which 
is  characteristic  of  the  sub-order,  Polysporea.     The  phenomenon  of  spore  formation 


MYXOSPORIDIA 


183 


is  not  simple  (fig.  97),  and  the  spore  itself  is  surrounded  by  a  bivalved  shell  or  sporo- 
cyst  and  contains  polar  capsules  in  addition  to  the  amoeboid  germ  (fig.  97,  G,  h).  The 
valves  of  the  sporocyst  and  the  polar  capsules  are  really  differentiated  nucleate  cells, 
so  that  each  spore  is  an  aggregate  of  cells  rather  than  one  cell,  though  only  a  single 
amoebula  issues  from  a  spore.  The  accounis  of  spore  formation  vary  somewhat 
according  to  the  different  workers. 

Spore  formation  is  usually  very  complicated  and  there  are  differences  of  opinion 
as  to  the  interpretation  of  various  stages,  pariictilarly  as  to  whether  conjugation 
occurs  therein.  The  process  is  initiated  by  the  concentration  of  cytoplasm  around 
one  of  the  nuclei  of  the  endoplasm,  so  that  a  small  spherical  mass  or  initial  corpuscle 
is  produced,  the  pansporoblast  (Gurley)  or  primitive  sphere  (Thelohan).  Sonje 
authors  state  that  a  pansporoblast  really  results  from  a  conjugation  of  two  ini'ial 
corpuscles  (fig.  97,  a— D).     Nuclear  multiplication  occurs  within  the  pansporoblast 


Fig.  97. — Myxobolus  pfeifferi.  Spore  formation.  A,  reproductive  cell  from  plasmodial 
trophoz"ite  ;  B,  cell  divided  unequally  into  two;  C,  smaller  cell  foiminjj  envelope  to  larger 
one  ;  D,  pinspornblast  formed  by  union  of  two  forms  like  C  ;  E,  multinucleate  pansporoblast 
two  of  the  nuclei  being  those  of  the  envelope  ;  F,  pinsporob'ast  divided  into  two  multinucleate 
sporoblasts  ;  G,  spore  differentiation  ;  /,  two  parietal  cells  forming  sporocyst  ;  be,  polar  cap- 
sules ;  a7n,  binucleate  amoebula  ;  11,  ripe  spore  in  which  the  two  nuclei  of  the  amoebula  have 
fused.     (After  Keysselitz.) 


(fig-  97?  e),  and  sooner  or  later  two  multinucleate  sporoblasts  are  formed  within  it 
(fig.  97,  f).  Each  sporoblast  gives  rise  to  a  single  spore,  which  consists  of  a  sporo- 
cyst or  envelope  composed  of  two  valves  each  secreted  by  a  cell,  two  polar  capsules 
each  secreted  by  a  cell,  and  the  sporoplasni  or  amoebula  which  becomes  binucleate 
(fig.  97,  g).  During  the  process  of  spore  formation  (fig.  97)  various  vegetative  and 
reduction  nuclei  maybe  produced,  in  addition  to  those  which  are  essentially  involved 
in  spore  formation,  and  the  sporocyst  cells  may  be  developed  early. 

Each  spore  contains  two  (figs.  94,  95)  or  more  polar  capsules  which  are  clearly 
visible  in  the  fresh  condition.  Each  polar  capsule  is  a  hollow,  more  or  less  pear- 
shaped  body,  secreted  by  a  cell  and  having  a  well  defined  contour.  Within  it,  a 
long,  delicate,  elastic  filament,  the  polar  filament,  is  formed,  and  lies  spirally  coiled 
in  the  polar  capsule  until  just  before  the  emergence  of  the  amoebula  from  the  spore 
(fi&-  95)-     The  polar  filament  is  ejected,  probably  under  the  influence  of  the  digestive 


184  THE   ANIMAL   PARASITES   OF   MAN 

juice,  when  the  spore  reaches  a  new  host,  and  serves  to  anchor  the  spore  to  the 
tissue  with  which  it  is  in  contact,  and  thus  allow  of  the  emergence  of  the  amoebula 
in  a  situation  suitable  for  its  development.  The  polar  capsule  with  its  contained 
polar  filament  has  been  compared  with  the  stinging  cells  or  nematocysts  of  the 
Coelentera,  but  it  has  a  totally  different  function. 

The  spores  fulfil  the  purpose  of  effecting  transmission  to  other  hosts.  In- 
fection occurs  by  the  ingestion  of  the  parasites  per  os  after  their  escape  by  some 
means  from  their  host.  Thelohan  and  others  have  demonstrated  that  the  valves  of 
the  spores  soon  open  under  the  influence  of  the  digestive  juices,  thus  allowing  the 
young  myxosporidia  to  escape.  Their  further  history  is  unknown  ;  but  it  may  be 
surmised  that  they  either  travel  direct  to  the  organs  usually  affected  (gall  bladder, 
urinary  bladder),  or  are  distributed  in  the  body  by  means  of  the  circulatory  or 
lymphatic  systems. 

The  Myxosporidia  that  invade  tissues  are  often  deadly  to  their  hosts.  They  may 
be  present  in  a  state  of  "  diffuse  infiltration  "  when  practically  every  organ  of  the 
body  may  be  infected,  as  in  barbel  disease  (due  to  Myxobolus  pfeifferi).  On  the 
other  hand,  the  parasites  may  be  concentrated  at  one  spot,  when  cysts,  either  large 
or  small,  are  produced.  Such  cysts  occur  on  the  gills  of  many  fishes.  A  few  addi- 
tional important  pathogenic  forms  are  Myxobolus  cyprini^  the  excitant  of  "  pocken- 
krankheit"  of  carp,  and  Lentospora  cerebralis,  parasitic  in  the  skeleton  of  Salmonidae 
and  Gadidas.  The  skeletons  of  the  tail,  fins  and  skull  particularly  are  seats  of  infec- 
tion, and  from  the  skull  the  Lentospora  can  spread  to  the  semicircular  canals,  result- 
ing in  loss  of  power  to  maintain  its  balance  on  the  part  of  the  fish.  On  this  account 
the  malady  is  termed  "drehkrankheit,"  Young  fish  are  more  particularly  infected. 
Myxobolus  7ieurobius  infects  the  spinal  cord  and  nerves  of  trout. 

Myxosporidia  are  divided  into  two  sub-orders — Disporea  and  Polysporea — 
according  to  whether  they  form  only  two  or  several  spores  during  their  growth. 
The  former  include  two  genera  limited  to  fishes,  which  are  easily  distinguishable  by 
the  shape  of  the  spores  :  Leptoiheca,  Thel.,  with  a  rounded  spore,  and  Ceratojnyxa^ 
Th^l.,  with  a  very  elongate  spore.  The  larger  number  of  genera  belong  to  the 
Polysporea,  which  are  divided  into  three  families : 

.    .  {' 

stam  wuh  iodine.  v  {b)  With  four  polar  capsules. — Chloromyxida. 

(2)  Amoeboid  germ  with  a  vacuole  stainable  with  iodine.     Spores  with  two    polar 
capsules. — Myxobolidcr. 
For  further  subdivisions  the  differences  in  the  spores  are  principally  utilized. 

Order.     Microsporidia.     Balbiani. 

These  are  the  organisms  discovered  in  the  stickleback  by  Gluge  in  1834,  and 
in  Coccus  hesperidum  by  Leydig  in  1853.  They  have  since  been  found  in  numerous 
other  arthropods,  especially  insects.  They  acquired  particular  importance  when  it 
was  discovered  that  they  were  the  cause  of  the  "pebrine"  disease  ("gattina"  of 
the  Italians)  which  caused  so  much  destruction  amongst  silkworms  {Bombyx  tnori). 
Pasteur  (1867-70)  and  especially  Balbiani  (1866)  participated  in  the  researches  on 
Nosema  botnbycis,  and  it  was  the  latter  who  classed  the  "pebrine  bodies"  or  "  psoro- 
spermia  of  the  arthropoda"  amongst  the  Sporozoaas  Microsporidia  (1882).^  The  com- 
plete life  cycle  of  N.  bonibycis  was  described  in  1909  by  Stempell.  The  Microsporidia 
are  not  confined  to  insects  and  arachnoids,  they  are  now  known  to  occur  also  in 
Crustacea,  worms,  bryozoa,  fishes,  amphibians  and  reptiles.  Certain  tumours  in 
fishes,  similar  to  those  formed  by  many  Myxosporidia,  are  produced  by  Micro- 
sporidia.    Fantham  and  Porter  found  that    Nosema  apis  was  pathogenic  to  bees 

'  C.  R.  Acad.  Set.,  Paris,  xcv,  p.  1168. 


(i)  Amoeboid  germ  with  a  vacuole  ["(«)  With  two  polar  capsules — Myxidiidcp. 
the  contents  of  which  do  not 


MICROSPORIDIA 


l8s 


and  other  insects,  and  was  the  causal  agent  of  the  so-called  "  Isle  of  Wight" 
disease  in  bees^  in  Great  Britain. 

The  Microsporidia,  as  their  name  implies,  form  minute  spores  which  usually  are 
oval  or  pear-shaped.  Each  spore  contains  a  single  polar  capsule  which  is  not  easily 
visible  in  the  fresh  state  (fig.  98,/)  and  a  single  amoeboid  germ  issues  from  the  sporfe 
(fig.  99.  b). 

The  life  cycle  of  Nosema  apis,  parasitic  in  bees,  may  be  taken  as  an  example  of 
that  of  a  microsporidian.  The  infection  of  the  host  is  initiated  by  the  ingestion  of 
spores  oi  N.  apis  in  food  or  drink  contaminated  with  the  excrement  of  other  infected 


Fig.  98. — Aosema  apis.  Various  .stages  in  life-cycle,  a,  planonts  or  amoebulse  from  chyle 
stomach  of  bee  ;  b,  amoeboid  planont  creeping  over  surface  of  gut  epthelial  cell ;  c,  uninucleate 
trophozoite  within  epithelial  cell  ;  d,  meront  with  nucleus  divided  into  four,  about  to  form 
four  spores  ;  e,  epiihelial  cell  cr(-wded  with  spores  ;  /",  young  spore  ;  g^  spore  showing  five 
nuclei,  polar  filament  ejected,  and  amoebula,  about  to  issue.  X  1,500,  a— ^;  X  2,150,/—^. 
(After  Fanlham  and  Porter.) 

bees.  Under  the  influence  of  the  digestive  juice  of  the  bee  the  spore-coat  (sporocyst) 
softens,  the  polar  filament  is  ejected  and  anchors  the  spore  to  the  gut  epithelium, 
and  the  minute  amoebula  contained  in  the  spore  emerges.  The  amoebula  is  capable 
of  active  amoeboid  movements  (fig.  98,  b)  and  so  is  termed  the  planont  or  wandering 
form  (fig.  98,  a).  After  a  short  time  each  planont  penetrates  between  or  into  the  cells 
of  the  epithelium  of  the  gut,  a  few  only  passing  through  into  the  body  cavity.  Within 
the  cells  the  amoebulce  become  more  or  less  rounded,  lose  their  power  of  movement, 
and  after  a  period  of  growth  of  the  trophozoite  (fig.  98,  c)  commence  to  divide  actively, 
these  dividing  forms  being  known  as  meronts  (fig.  98,  d).  Various  forms  of  fission 
occur,  and  during  this  phase,  termed  merogony,  the  numbers  of  the  parasite  within 
the  host  are  greatly  increased,  with  concomitant  destruction  of  the  epithelium 
{fig.  98,  e).  After  a  time  sporogony  cominences.  The  full-grown  meront  becomes 
successively  the  pansporoblast  and  sporoblast.  Nuclear  multiplication  and  differen- 
tiation ensue  and  five  nuclei  are  ultiinately  produced.  At  the  same  time  a  sporocyst 
is  secreted,  and  two  vacuoles  are  produced  within.     One  is  the  polar  capsule,  and 


*  Annals  Trop.  Med.  and  Parasitol.,  vi,  pp.  145-214,  3  pis. 


1 86 


THE   ANIMAL   PARASITES   OF   MAN 


within  it  the  polar  filament  is  differentiated  ;  the  other  forms  the  posterior  vacuole 
(fig-  98,  ^).  Between  the  two  vacuoles  the  body  cytoplasm  or  sporoplasm  forms  a 
girdle-like  mass.  Of  the  nuclei,  one  regulates  the  polar  capsule,  two  control  the 
secretion  of  the  sporocyst,  and  two  remain  in  the  sporop'asm.  The  polar  capsule  and 
polar  filament  are  not  usually  visible  in  the  fresh  condition,  but  can  be  demonstrated 
by  the  use  of  various  chemical  reagents  (fig.  100).  The  sporoplasm  ultimately  becomes 
the  amoebula  (fig.  98,  g)  which  issues  from  the  spore  after  the  ejection  of  the  polar 
filament. 

A  trophozoite  (meront)  of  N.  apis  becomes  a  single  pansporoblast  which  gives 
rise  to  one  sporoblast  producing  one  spore,  and  this  procedure  is  characteristic  of 
the  genus  Noseina.  In  other  genera  the  trophozoite  may  form  more  than  one  pan- 
sporoblast and  each  pansporoblast  may  form  a  variable  number  of  spores  in  different 
cases.  Various  attempts  at  classification  have  been  based  on  these  characteristics. 
It  must  suffice  here  to  note  that  in  the  cases  where  the  trophozoite  becomes  one 
pansporoblast,  the  latter  can  produce  four  spores  in   the  genus  Gurleya,  eight  spores- 


Fig.  99. — a,  section  through  the  abdominal 
wall  of  a  silkworm,  whose  epithelial  ceils 
contain  Microsporidia  {Nosenia  bombycis)  ;  b, 
a  spore,  the  contents  of  which  are  escaping. 
(After  Balbiani.) 


Fig.  100. — Nosenia  bombycis,  Naeg.. 
Spores  treated  with  nitric  acid,  thus 
rendering  the  polar  capsule  perceptible, 
and  the  filament  has  protruded  from  one 
of  the  spores.     (After  Thelohan.) 


in  Thelohania  and  many  spores  in  Pleistophora.  In  other  cases,  where  the  tropho- 
zoites give  rise  to  many  pansporoblasts,  each  of  the  latter  may  form  many  spores,, 
as  in  the  genus  Glugea. 

A  few  pathogenic  microsporidian  parasites  other  than  N.  apis  may  be  mentioned. 
N.  bombycis^  causing  pebrine  in  silkworins,  may  infect  any  or  all  the  tissues  of  the 
host  (fig.  99).  The  larvie  of  the  host,  i.e.^  the  "  silkworms,"  may  become  infected  by 
eating  food  contaminated  with  spore-containing  excrement  of  already  infected  silk- 
worms. In  cases  of  heavy  infection  the  silkworm  dies,  but  should  the  infection  be 
less  intense  the  larva  becomes  a  pupa  in  which  the  parasite  persists,  so  that  the  moth 
emerges  from  the  cocoon  already  infected.  Not  only  is  the  moth  parasitized  itself,, 
but  the  Nosema  reaches  the  generative  organs  of  both  sexes  and  penetrates  the 
ovaries  of  the  female,  with  the  result  that  the  ova  are  deposited  infected.  Such 
infected  eggs  are  capable  of  developing,  so  that  infection  may  be  transmitted 
hereditarily  as  well  as  by  the  contaminative  method.  Infected  eggs  can  be  recog- 
nized by  microscopic  examination,  as  Pasteur  showed,  and  thus  preventive  measures^ 
may  be  adopted. 

A  microsporidian  parasite  is  known  to  occur  on  the  roots  of  the  spinal  and  cranial 
nerves  of  Lophius  piscatorius^  the  angler  fish.  This  parasite  is  variously  referred  to 
the  genera  Nosema  and  Glugea. 

Thelohania  contejeani^  parasitic  in  the  muscles  of  crayfish,  is  believed  by  some 
to  be  the  causal  agent  of  recent  epizootics  among  them,  though  others  believe  the 
disease  to  be  really  due  to  a  bacillus.  It  may  be  that  the  one  organism  aids  in  the 
entry  of  the  other  into  the  host. 


ACTINOMYXIDIA 


187 


Order.     Actinomyxidia,  Stole. 

A  brief  mention  may  be  made  of  the  Actinomyxidia  (fig.  loi),  which  were  first 
described   by   Stole   in    1899  as    parasites  of  Oligochaetes.     They  have   also  been 
investigated   by  Mrazek,   and   a   detailed   study  of  certain   species  was  made   by 
Caullery  and  Mesnil  (1905).     The  tropho- 
zoite is  small  and  amceboid.     The  spores 
are  large,  and  exhibit  tri-radiate  symmetry. 
Spore  formation  is  complicated  and  sexual 
processes  occur  therein.     Many  amoebulse 
are  set  free  from  each  spore. 

Order.      Sarcosporidia,  Balbiani. 

The  first  member  of  this  group  was 
discovered  by  Miescher  in  1843.  '^1^'s 
author  found  white  filaments  running 
parallel  with  the  direction  of  the  fibres 
in  the  voluntary  muscles  of  mice.  They 
were  visible  to  the  naked  eye,  and  proved 
to  be  cylindrical  tubes  tapering  at  each 
end.  They  were  as  long  as  the  muscular 
fibres,  were  enveloped  in  a  membrane,  and 
contained  innumerable  elongate  or  kid- 
ney-shaped bodies  and  a  smaller  number 
of  little  spherical  forms.  Th.  v.  Hessling 
confirmed  (1853)  the  occurrence  of  these 
"  Miescher's  tubes  "  within  the  muscular 
fibres,  this  author  having  discovered  the 
same  structures  in  the  heart  muscles  of 

deer,  cattle,  and  sheep.  Both  investigators  considered  them  to  be  pathological  trans- 
formations of  the  muscles,  v.  Siebold,  from  his  own  experiences,  regarded  them 
as  fungus-like  entophytes. 

Rainey  (1858)  discovered  similar  structures  in  the  muscular  system  of  pigs,  and 
considered  them  to  be  early  stages  of  Cysticercus  cellulosce,  which  error  Leuckart 
rectified,  simultaneously  emphasizing  their  relationship  with  Myxosporidia.  Both 
these  authors  found  them  in  the  muscular  fibres,  and  both  observed  that  they 
possessed  a  thick  striated  membrane.  Manz  (1867)  published  the  results  of  more 
minute  investigations  on  the  structure  and  contents  of  the  cylinders.  This  observer 
also  recognized  the  disease  in  rabbits  and  attempted  to  cultivate  the  parasites.  He 
also  tried  to  induce  experimental  infection  in  guinea-pigs,  rats,  and  mice,  but  the 
result  was  negative. 

However,  domestic  and  wild  mammals  are  not  the  only  hosts  of  Sarcosporidia  ; 
these  parasites  are  also  harboured  by  birds.  Thus,  according  to  Kiihn,  they  are 
found  in  the  domestic  fowl  ;  according  to  Rivolta  in  Tiirdiis,  Corvtis^  and  other 
birds  ;  according  to  Stiles  in  North  American  birds  ;  while  Fantham  found  Sarco- 
sporidia in  the  African  mouse-bird.  Coitus.  Reptiles  also  are  parasitized  occasionally. 
Bertram  found  them  in  the  gecko,  Liihe  in  the  wall-lizard.  It  was  found  also  that 
the  Sarcosporidia  could  develop  not  only  in  the  muscles  but  also  in  the  connective 
tissue.  This  led  to  the  foundation  of  a  new,  but  provisional,  classification  by  Blan- 
chard,  using  the  generic  name  Miescheria  for  the  parasites  in  the  muscles  and 
Balbiania  for  those  in  the  connective  tissue.  Finally,  Sarcosporidia  have  also  been 
observed  in  man. 


Fig.  10  r. — Spore  of  Hexactinomyxon 
psamnioryctis.  At  top  of  .figure  three  polar 
capsules,  one  with  polar  filament  extended. 
X  450.     (Aftei  Stole.) 


88 


THE   ANIMAL   PARASITES   OF   MAN 


The  relation  of  these  parasites  to  certain  diseases  of  domestic  animals  has  been 
studied  by  veterinary  surgeons.  Sarcosporidia  may  cause  fatal  epizootics  among 
sheep. 

There  is  still  a  wide  field  open  for  research  in  regard  to  the  structure  and  develop- 
ment of  these  parasites,  and  the  manner  in  which  the  hosts  become  infected. 

The  Sarcosporidia  usually  appear  as  elongate,  cylindrical,  or 
fusiform  bodies,  rounded  at  both  extremities  and  of  various  lengths 
and  breadths  (fig.  102).  In  some  species  they  may  be  from  16  mm. 
to  50  mm.  long,  as  in  the  sheep  and  roebuck.  These  bodies  are  the 
so-called  sarcocysts  or  Miescher's  tubes.      They   He    in   transversely 


Fig.  102. — Longitudinal  section  of  a 
muscle  of  the  domestic  pig,  with  Sar- 
cocystis  niiescheriana.  x  30.  (After 
Kiihn.) 


Fig.  103. — Transverse  section  of  the 
muscle  of  a  pig,  wiih  Sarcocystis 
niiescheriana.      x  38.     (After  Kiihn.) 


striated  muscular  fibres  which  they  distend  more  or  less.  The  forms 
found  in  the  connective  tissue  are  apparently  parasites  which  originally 
inhabited  the  muscular  fibres,  and  only  on  disintegration  of  the  fibres 
reached  the  connective  tissue,  where  they  grow  to  large  oval  or  globular 
bodies  (fig.  105).  The  mammalian  muscles  usually  infected  are  those 
of  the  oesophagus,  larynx,  diaphragm,  body-wall,  and  the  psoas  muscles. 
The  skeletal  muscles  may  be  affected  in  acute  cases,  as  well  as  those 
of  the  tongue  and  eye.     The  heart  muscles  are  sometimes  parasitized. 

In  fresh  material  cut  into  thin  slices  the  parasites  are  frequently  recognizable,  even 
with  the  naked  eye,  because  of  their  yellowish-white  colour.  Under  the  microscope 
they  appear  to  be  coarsely  granular  (fig.  103).  Beginners  may  find  some  difficulty 
in  distinguishing  them  from  other  foreign  bodies,  such  as  dead  and  calcified  encap- 
suled  Trichinae,  or  from  Cysticerci  that  have  died  and  become  calcified  in  the  early 
stages,  more  particularly  as  the  Sarcosporidia  also  occasionally  may  become  calcified. 

The  Sarcosporidia  are  always  enveloped  in  a  membrane,  which 
is  probably  formed  at  an  early  stage.     In  a  few^  cases  it  remains  thin 


SARCOSPORIDIA 


189 


and  simple,  in  other  cases  a  radially  striated  ectoplasmic  layer  is 
present  (figs.  104,  108),  which  has  been  variously  described.  From  the 
inner  integument,  which  may  be  homogeneous  or  fibrous,  thick  or 

thin,  membranes  or  trabeculae  pass  into  the 
interior  of  the  body,  forming  anastomosing 
partitions,  and  so  producing  a  system  of 
chambers  of  various  sizes  that  do  not  com- 
municate with  one  another  (figs.  104,  108). 
These  chambers  are  occupied  by  sickle-  or 
bean-shaped  bodies  (spores  or  sporozoites), 
or  various  developmental  stages  of  them. 
The  oldest  spores  are  found  in  the  centre  of 
the  Miescher's  tubes  or  trophozoites.  If  they 
are  not  liberated  they  die  there,  so  that  the 
central  chambers  of  the  tube  are  empty  and 
hollow. 

In  the  youngest  Sarcosporidia  (40  /x  in 
length)  from  the  muscles  of  the  sheep  there 
occur,  according  to  Bertram,  small  roundish 
or  oval  cells  (4^11  to  5 /x),  the  nuclei  of  which 
are  half  their  size,  and  are  embedded  in  a 
granular  protoplasmic  mass.  In  somewhat 
larger,  and  therefore  older,  cylinders,  the 
investing  membrane  of  which  already  shows 
both  layers,  the  cells  have  become  larger  (to 
7  fx)  and  are  more  sharply  outlined  from  each 
other  (fig.  106).  These  uninucleate  cells  may 
be  considered  as  pansporoblasts.  In  each 
pansporoblast  division  of  the  nucleus  occurs 
(fig.  107),  and  meanwhile  'the  pansporo- 
blasts become  isolated  within  the  chambers,  the  dividing  partitions 
of  which  originate  from  the  granular  protoplasm  which  is  present 
between  the  pansporoblasts.  The  numerous  uninucleate  daughter 
forms  produced  within  the  chambers  become  spores  direct  (fig.  108). 
The  process  commences  in  the  centre  of  the  cylinders  or  sarco- 
cysts,  and  then  progresses  towards  the  extremities,  the  parasites  mean- 
while increasing  in  size,  and  new  pansporoblasts  being  continually 
formed  at  the  extremities  (fig.  107). 

The  spores  (sometimes  called  Rainey's  corpuscles),  vary  in  shape 
according  to  the  species,  but  are  also  of  different  form  individually. 
They  are  mostly  kidney-,  bean-  or  sickle-shaped  (fig.  109),  and  of 
small  size,  sometimes  reaching  14  /x  by  3  yt*  to  5  /x.  They  are 
apparently  surrounded  by  a  thin  membrane,  and  at  one  extremity 
(according  to  the  discovery  of  L.  PfeifTer,  confirmed  by  van  Eecke, 


Fig.  10^, — Sai  cocys.isniies- 
cheriana  from  pig.  Late 
stage  in  which  body  is  divided 
into  numerous  chambers  or 
alveoli,  each  containing  many 
spores.  {From  Wasielevirski, 
after  Manz.) 


190 


THE   ANIMAL   PARASITES   OF   MAN 


/ 
/    • 


f. 


a 


Fig.  105. — Transverse  section  of  Sarcocystis  tenella^ 
Raill.  Fiom  the  oesophagus  of  the  sheep,  Ovis  aries. 
X  38.  a,  marginal  chambers  filled  with  spores  ;  d,  con- 
nective tissue  of  the  oesophagus ;  c,  muscles  of  the 
oesophagus. 


Fig.  106. — Young 
Sarcocystis  tenella  of 
the  sheep,  47  ^i. 
in  length.  (Alter 
Bertram.) 


Fig.  107.— End  of  a 
trophozoite  c.f  Sarcocystis 
miescheriana  fr  -m  the 
diaphr  gin  of  the  pig, 
showing  riivi-ion  in  pan- 
sport  bbss.  X  800. 
(After    Bertram.) 


Fig.  108.  —  Sarcocystis 
hlanchardi  of  the  ox.  Longi- 
tudinal section  of  sarc«»cy>t  or 
Miescher's  lube.  a,  sub- 
stance of  muscle  fibre  ;  b, 
envelope  of  saro'cysi ;  <:,  mus- 
cle nuclei  ;  r/,  sports  in  cham- 
bers ;  ^,  ground  subs'ai)ce. 
X  400.  (From  Wasielcwski, 
after  van  Eecke.) 


SARCOSPORIDIA  191 

Laveran  and  Mesnil)  contain  an  obliquely  striated  body  (fig.  109)  often 
homologized  with  the  polar  capsule,  while  the  greater  part  of  the  spore 
is  taken  up  by  the  nucleate  sporozoite.  Several  authors  state  that  they 
have  also  observed  filamentous  appendages  (polar  filaments)  at  one 
end  of  the  spores,  and  have  seen  two  kinds  of  spores  in  the  same 
Sarcosporidium.  Spores  of  various  species  of  Sarcosporidia  may 
contain  metachromatic  granules,  often  centrally  placed  (fig.  109). 
These  granules  may  be  metabolic  or  possibly  may  contain  toxin 
(see  below). 

The  gymnospores  of  Sairocystis  jiiuris,  fr  om  the  mouse,  show  active 
boring  movements  when  kept  in  saline  solution  warmed  to  35°  or 
37°  C.  S.  inuris  is  very  deadly  to  its  host.  From  their  structure  the 
spores  do  not  appear  to  have  great  powers  of  resistance  to  external 
conditions.     They  measure  1 2  yu,  by  3  ytt  to  4  />«-  or  less. 

Laveran  and  Mesnil  (1899)  isolated  a  toxin  from  S.  tenella  of 
the  sheep  and  called  it  sarcocystin.  This  substance  is  especially 
pathogenic  to  experimental  rabbits. 

The  duration  of  life  of  the  Sarcosporidia  is  a  comparatively  long 
one.  The  affected  muscular  fibres  may  remain  intact  and  capable  of 
performing  their  functions  for  a  long  time, 
but  at  last  they  perish,  if  the  host  lives  long 
enough.  Thus  the  Sarcosporidia  of  the 
muscles  are  then  enveloped  only  by  sarco- 
lemma,  and  finally,  when  this  likewise  dis- 
appears, they  fall  into  the  intra-muscular 
connective   tissue.  In  many  cases  the  Sarco-  ^  ^ 

sporidia  die  off  within  their  hosts,  this,  Fig.  109.— Spores  of  .Sar^^?- 
according  to  Bertram,  being  brought  about  ^otnTt'  H-  c^a.S; 
by  a  disintegration  of  the  spores  in  the  cen-  b,  stained,  showing  meta- 
tral  chambers.  In  other  cases  the  leucocytes  ^f,^'''^'^  ^'^,^^;'.  ""(Vt"; 
play  a  part  in  the  destruction  of  the  Sarco-  Laveran  and  Mesnil.) 
sporidia,  and  sometimes  it  happens  that  lime 
salts  are  deposited  in  and  around  the  vacant  cylinders. 

In  some  places  pigs,  sheep,  mice  and  rats  are  infected  with 
sarcosporidiosis  to  a  remarkable  extent,  in  certain  cases  almost  reach- 
ing 100  per  cent.  Young  animals  also  are  infected,  and  perhaps 
infection  only  takes  place  during  youth. 

Although  the  natural  mode  of  transmission  of  the  Sarcosporidia 
remains  to  be  determined,  yet  various  experimental  researches  on 
the  problem  are  of  interest  and  importance.  Theobald  Smith  (1901) 
found  that  mice  could  be  experimentally  infected  with  S.  miiris 
by  feeding  them  with  the  flesh  of  other  infected  mice.  The  incuba- 
tion period  was  a  long  one,  namely  forty  to  fifty  days.  Thus,  on  the 
forty-hfth  day  after  feeding  young  Sarcosporidia  were  found,  and 
seventy  days  after  feeding  spore  formation  began.  Ripe  spores  were 
found  two  and  a  half  to  three  months  after  the  commencement  of 


192  THE   ANIMAL   PARASITES   OF    MAN 

these  experiments.  This  mode  of  infection — a  cannibahstic  one — 
hardly  seems  likely  to  be  the  natural  method  for  the  infection  of  sheep 
and  ruminants  generally.  Smith's  researches  have  been  confirmed. 
Negre'  (1910)  found  that  the  faeces  of  mice  fed  on  infected  muscular 
tissue  were  infective  to  other  mice  when  ingested  by  them.  Negri*^ 
infected  guinea-pigs  with  S.  nuirls  by  feeding  them  on  infected  mouse 
flesh,  and  found  that  the  parasite  in  guinea-pigs  showed  different 
characters  from  those  exhibited  by  it  in  mice.  Darling^  also  succeeded 
in  infecting  guinea-pigs  with  S.  mnrisy  and  Erdmann  infected  mice 
with  S.  tenella  (from  the  sheep). 

According  to  Erdmann*  (iqio)  the  Sarcosporidian  spore  germinates 
in  the  intestine  of  the  host,  which  has  recently  ingested  infected 
material.  The  spore  liberates  its  contained  toxin  —  sarcocystin  — 
which  acts  upon  the  adjacent  intestinal  epithelium,  whereby  the  latter 
is  shed,  and  an  amcebula  creeps  out  of  the  spore.  The  amoebula  is 
able  to  penetrate  the  denuded  area  and  get  directly  into  the  lymph- 
spaces  of  the  submucous  coat  of  the  intestine.  The  first  period  of 
development,  lasting  some  twenty-eight  to  thirty  days,  is  said  to  be 
passed  in  the  lymph-spaces  of  the  intestine.  Later  the  amoebula 
reaches  a  muscle  fibre.  Writing  in  May,  1914,  Erdmann^  records  the 
appearance  of  small  amoeboid  and  schizogony  forms  six  days  after 
infection  of  the  host.  Crawley^  (19 13)  controverts  some  of  these 
statements  and  considers  that  the  Sarcosporidian  spore,  still  sickle- 
shaped,  bores  its  way  into  the  epithelial  cells  of  the  intestine  and 
comes  to  rest  there.  The  spore  then  becomes  round  or  elliptical,  and 
peripheral  masses  of  chromatin  appear  within  it,  suggesting  schizogony. 
This  happens  about  twelve  hours  after  feeding,  and  in  twenty-four 
hours  the  spores  appear  to  have  left  the  intestine.  More  recently  (May, 
1914),  Crawley^  considers  that  there  is  sexual  differentiation  among  the 
Sarcosporidian  spores,  a  few  hours  after  their  ingestion  by  the  host. 

Interesting  discussions  have  occurred  as  to  the  site  of  the  toxic 
sarcocystin  within  the  spore.  Metachromatic  granules  occur  in  the 
middle  of  the  Sarcosporidian  spore  (fig.  109),  and  the  toxin  may  be 
contained  in  these  grains,  as  they  disappear,  according  to  Erdmann, 
before  the  amoebula  penetrates  the  denuded  intestinal  wall.  However, 
a  polar  capsule,  containing  a  polar  filament,  may  be  present  at  one  end 
of  a  Sarcosporidian  spore.  Laveran  and  Mesnil  described  a  striated 
area  at  the  more  pointed  end  of  the  spore  of  S.  tenella^  which 
area  tliey  consider  to  represent  a  polar   capsule.     Fantham^  (1913) 

'   C.  R.  Soc.  Biol.,  Ixviii,  p.  997. 

'^  Cefztralbl.  f.  Bakt.,  Orig.,  xlvii,  p.  612  ;  see  also  xlvii,  p.  56  ;   Iv,  p.  373. 

^Joitrn.  Exptl,  Med.,  xii,  p.  19.  ■»  Sitz.  Gesell.  naturf.  Freunde  zti  Berlin,  p,  y]-]. 

^Proc.  Soc.  Exper.  Biol,  and  Med.,  xi,  p.  152.  ^Science,  xxxvii,  p.  498. 

^  Proc.  Acad.  Nat.  Sci.,  Philadelphia,  May,  1914,  p.  432. 

^  Proc.  Cambr.  Philosoph.  Soc.y  xvii,  p.  221. 


SARCOSPORIDIA  I93 

found  a  vacuole-like,  polar  capsule  area  in  the  spores  of  S.  colli 
from  the  African  mouse-bird.  The  sarcocystin  may  be  contained 
in  the  polar  capsule.  The  nucleus  of  the  spore  is  generally  at  the 
opposite,  blunter  end. 

Again,  various  authors  have  stated  that  Sarcosporidian  spores  may 
occur  in  the  blood  of  the  host  at  times.  If  so,  then  an  intermediate 
host  may  be  concerned  in  their  transmission.  Perrin  suggested  that 
Sarcosporidia  might  be  spread  by  blow-flies  and  flesh-flies. 

The  classification  of  the  Sarcosporidia  as  proposed  by  R. 
Blanchard,  which  was  based  on  their  various  habitats,  can  no  longer 
hold,  because  the  same  species  may  occur  in  the  muscles  as  well  as 
in  the  connective  tissue.  For  the  present,  the  few  species  that  are 
known  maybe  placed  in  one  genus,  Sarcocystis,  Ray  Lankester,  1882. 

The  following  species  of  Sarcocystis  are  of  interest  : — 

S.  iniescheriana,  Kiihn,  1865,  in  the  pig. 

S.  bertrami,  Doflein,  iQoi,  in  the  horse. 

S.  teiiella,  Railliet,  1886,  in  sheep.  S.  tenella  bubali  in  buffaloes  in 
Ceylon  and  Egypt. 

S.  blanchardi,  Doflein,  1901,  in  cattle. 

S.  niuris,  Blanchard,  1885,  in  the  mouse,  to  which  it  is  lethal. 

S.  Iiiieti,  Blanchard,  1885,  in  the  seal. 

S.  coin,  Fantham,  191 3,  in  the  African  mouse-bird,  Coliiis 
eryt/ironieloii. 

Also  various  Sarcosporidia  from  antelopes,  monkeys,  opossum, 
birds,  the  gecko  and  wall-lizard  are  known. 

The  spores  of  S,  mnris,  S.  bertrami,  S.  tenella,  and  S.  colli  can 
multiply  by  longitudinal  fission. 

Sarcosporidia  observed  in  Man. 

(i)  Lindemann  ^  found  on  the  valves  and  in  the  myocardium  of 
a  person  who  had  died  of  dropsy  certain  brownish  masses,  3  mm.  in 
length  and  1-5  mm.  in  breadth  which  he  regarded  as  gregarines.  If 
these  were  actually  independent  animal  organisms  it  may  be  suggested 
that  they  were  Sarcosporidia.  Rivolta  (1878)  named  the  species 
S.  liiidcjuanni. 

(2)  Rosenberg^  found  a  cyst  5  m_m.  in  length  and  2  mm.  in  breadth 
in  a  papillary  muscle  of  the  mitral  valve  of  a  woman,  aged  40, 
who  had  died  from  pleuritis  and  endocarditis.  The  cyst  contained 
no  scolex  nor  booklets  of  taenia.  Numerous  small  refracting  bodies, 
round,  oval  or  kidney-shaped,  were  found  in  a  daughter  cyst,  as  well 

'  **  Ueb.  d.  hyg.  Bdtg.  d.  Gregarinen,"  Dtsche.  Ztschr.  f.  Staatsarzneikunde,  1868,  xxvi, 
p.  326. 

*  "  Ein  Befund  von  Psorosp.  in  Herzmusk  d.   Menschen,"  Ztschr.  f.  Hygiene,  1892,  xi, 

P-  435- 
^3 


194  THE    ANIMAL    PARASITES    OF   MAN 

as  sickle-shaped  bodies.     The  description  hardly  appears  to  indicate 
Sarcosporidia. 

(3)  Kartulis^  observed  Miescher's  cylinders  of  various  sizes  in 
the  liver  (?)  and  in  the  muscular  system,  of  a  Sudanese  who  had 
succumbed  to  multiple  abscesses  of  the  liver  and  abdominal  muscles. 
This  may  be  considered  as  the  first  actual  case  of  the  occurrence  of 
Sarcosporidia  in  man.     Koch  in  1887  described  a  case  in  Egypt. 

(4)  The  case  reported  by  Baraban  and  St.  Remy^  was  at  once 
demonstrated  as  certain.  It  related  to  a  man  who  had  been  executed, 
and  in  the  laryngeal  muscles  of  whom  Sarcosporidia  were  found  ;  the 
length  of  the  parasites  varied  between  150  //,  and  1,600  /a,  their 
breadth  between  77  /x  and  168  ^.  The  affected  muscular  fibres  were 
distended  to  four  times  their  normal  thickness.  This  species  was 
described  by  Blanchard  as  *^  Miescheria "  inuris,  but  according  to 
Vuillemin,  it  was  more  probably  Sarcocystis  tenella  of  the  sheep. 

(5)  Vuillemin  has  also  described  a  case  of  Sarcosporidia  found 
in  the  muscles  of  a  man  who  died  from  tubercle  at  Nancy.  The 
author  considered  that  the  parasite  corresponded  to  S.  tenella. 

(6)  Darling^  (1909)  found  Sarcosporidia  in  the  biceps  of  a  negro 
from  Barbados. 

The  Myxosporidia,  Microsporidia,  Actinomyxidia  and  possibly  the 
Sarcosporidia  may  be  included  within  the  section  Cnidosporidia 
(Doflein),  since  they  possess  spores  containing  polar  capsules. 

Order.     Haplosporidia,  Caullery  and  Mesnil. 

The  Haplosporidia  are  a  group  of  organisms  having  both  a  simple  structure  and 
life-history.  The  simplicity  may  represent  a  primitive  condition  or  may  be  due  to 
degradation  resultant  on  parasitism,  and  thus  it  is  possible  that  the  group  is  not 
a  homogeneous  one.  The  order  Haplosporidia  was  created  by  Caullery  and  Mesnil 
in  1899,  and  includes  parasites  of  rotiifers,  annelids  (fig.  no),  Crustacea,  fish,  pro- 
chordates  and  man.  They  may  be  present  in  the  body  cavity  or  alimentary  tract, 
and  can  also  occur  in  the  septum  nasi  of  man,  in  the  nervous  system  of  Cephalo- 
discus,  and  in  tumours  of  fish. 

As  the  name  implies,  the  spores  of  the  Haplosporidia  are  simple,  without  polar 
capsules,  and  are  uninucleate.  In  some  genera,  e.^.,  Haplosportdium,  Urospori- 
dium  (fig.  Ill)  there  is  a  spore-coat  or  sporocyst  which  may  be  elongate  or  spiny. 
The  developmental  cycle  of  a  Haplosporidian,  such  as  Haplosporidium  or  Bertramia, 
begins  with  a  small,  uninucleate  cell,  often  rounded,  possessing  a  cell  membrane  that 
may  be  prolonged  into  processes.  Growth  takes  place,  coupled  with  an  increase  in 
the  number  of  nuclei,  so  that  a  multinucleate  trophozoite  is  produced.     Later,  this 


'  Kartulis,  *'Ueb.  pathog.  Protoz.  b.  Menschen,"  Ztschr.  f.  Hyg.  u.  Inf.,  1893,  xiii,  p.  1. 
■Compare  also  Braun,  M.,  Die  Thier.  Par.  d.  Mensch.,  2nd  Edit.,  Wrzbg.,  1895,  p.  92  ; 
Braun,  M.,  "Z.  York.  d.  Sarcosp.,  b.  Menschen,"  Centralbl.  f.  Bakt.  1895.  xviii,  p.  13. 

'"  Sur  un  cas  de  Tub.  Psorosp.  ob.  chez  rhomme,"  C.  R.  Soc.  Biol.,  Paris,  1894  (x),  I, 
p.  201.     "  Le  Parasitisme  d.  Sarcosp.  chez  rhomme,"  Bibliogr.  Anat.  1894,  p.  79. 

^  Arch.  Internal  Med.,  Ill,  p.  183. 


HAPLOSPORIDIA 


195 


multinucleate  trophozoite  becomes  segmented  into  a  number  of  ovoid  or  spherical 
pansporoblasts,  which  give  rise  to  few  (one  to  four)  spores.  Such  a  spore,  when  set 
free,  begins  the  life  cycle  over  again. 

More  recently  (1905- 1907)  two  important  organisms  have  been  described  and 
included  in  this  group,  namely,  Neurosporidium  cephalodiscP  (Ridewood  and 
Fantham)  from  the  nervous  system  of  the  prochordate,  Cephalodisciis  nigrescens, 
and  Rhi7iosporidiu7n  kinealyi  (or  seeberi)  from  the  septutn  nasi  of  man.  In  the  case 
<ii  Rhino sporidiiim  and  Neiirosporidiitin^  after  the  uninucleate  spore  has  grown  into 


f 


5 


Fig.  1 10.  — Haplosporidiuni  heterocirri. 
Section  throughout  wall  of  the  Polychtete 
worm,  Heterocirrtis  viridis^  showing  various 
developmental  stages  of  the  Haplosporidiuin. 
X  550.     (After  Caullery  and  Mesnil.) 


Fig.  III. — Haplosporidian  spores, 
a.  /;,  Haplosporidhini  heterocirri.  a, 
fresh  ;  b,  after  immersion  in  sea  water  ; 
Cy  d,  Urosporidium  ftiliginosum. 
X  1000.    (After  Caullery  and  Mesnil.) 


a  multinucleate  trophozoite,  the  latter  segments  into  uninucleate  pansporoblasts, 
^as  in  the  preceding  cases.  A  difference  then  occurs,  for  each  pansporoblast 
-enlarges,  its  nucleus  divides  and  a  "  spore-morula"  is  formed.  Thus  a  multinucleate 
pansporoblast  or  spore-morula,  divided  into  many  uninucleate  sporoblasts  (spore 
mother  cells)  is  produced,  and  each  sporoblast  without  further  change  becomes 
a,  iminucleate  spore. 

The  Haplosporidia  have  therefore  been  divided  by  Ridewood  and  Fantham  (1907)  ^ 
into  two  sections  : — 

(i)  The  Polysporulea,  wherein  the  pansporoblast  gives  rise  to  a  number  of 
spores  (nine  or  more),  e.g.,  Rhinosporidium,  Neurosporidium. 

(2)  The  Oligosporulea,  wherein  the  pansporoblasts  give  rise  each  to  a  few  (four) 
spores  or  to  only  a  single  spore,  e.g.,  Haplosporiditan,  Bertrainia,  Ccelosporidium, 
Ichihyosporidiiwi. 

Rhinosporidium  kinealyi,  Minchin  and  Fantham,  1905. 

Rhinosporidhnn  kinealyi,  parasitic  in  man,  must  now  be  con- 
sidered in  greater  detail.  This  organism  was  found  in  nasal  polypus 
in  India,  and  has  since  been  recorded  from  the  ear  as  small  nodules 
in  the  external  auditory  meatus.     The  Indian  cases  came  from  the 


^    Quart.  Journ.  Microsc.  Sci.,  li,  p.  81. 

-  See  Fantham,  Brit.  Assoc.  Reports,  1907,  p.  553. 


196  THE   ANIMAL   PARASITES   OF   MAN 

neighbourhoods  of  Calcutta  and  Madras,  and  the  parasite  has  been 
seen  in  Ceylon.  Similar  structures  have  since  been  described  from 
the  United  States  and  South  America. 

The  Rhinosporidium  polypus  is  said  not  to  be  particularly  painful^ 
though  nasal  forms  must  interfere  with  breathing  to  some  extent.  The 
first  nasal  polyp  reported  from  India  formed  a  vascular  pedunculated 
growth  on  the  septum  nasi  and  was  about  the  size  of  a  large  pea  or 
raspberry.  It  was  compared  with  a  raspberry,  being  red  in  colour 
with  a  number  of  small  whitish  dots  upon  its  surface.  When  the 
tumour  was  cut,  a  number  of  similar  whitish  dots  were  seen  within. 
These  were  the  cysts  of  Rhinosporidium.  According  to  Minchin 
and  Fantham^  (1905);  they  vary  considerably  in  size  and  measure  up 
to  200  yu-  or  250 /x  in  diameter.  Each  possesses  a  cyst  wall  which  varies 
in  thickness  in  different  cysts.  Its  outer  wall  is  always  firm  and 
distinct,  the  inner  limit  being  less  definite  at  times.  Each  large  cyst 
is  filled  with  numbers  of  spherical  or  oval  bodies,  showing  every 
gradation  between  small  ones  at  the  periphery  and  large  ones  at  the 
centre  (fig.  112).  Roughly,  three  zones  of  parasites  can  be  dis- 
tinguished in  a  large  cyst,  a  peripheral  set  consisting  of  the  youngest 
parasites,  an  intermediate  group  and  a  central,  oldest  zone.  A  large 
cyst  may  possess  a  pore  for  the  egress  of  its  contents.  Some  of  the 
cysts  show  polar  distribution  of  the  zones. 

The  youngest  forms  of  Rhinosporidium  are  difficult  to  detect. 
They  are  small,  granular  masses,  round,  ovoid  or  irregular  and  at  times 
even  amoeboid  in  appearance.  These  are  young  trophozoites.  They 
increase  in  size,  but  encystment  occurs  early,  the  outer  layer  becoming 
firm  so  that  the  organisms  have  a  definite  contour.  Each  is  soon 
multinucleate  and  the  cytoplasm  segments  around  the  nuclei.  The 
cyst  thus  becomes  full  of  uninucleate  pansporoblasts  or  sporonts, 
with  a  peripheral  layer  of  undifferentiated  protoplasm.  The  pan- 
sporoblasts grow  in  size.  In  the  larger  cysts  the  formation  of 
pansporoblasts  progresses  at  the  expense  of  the  peripheral  layer  of 
protoplasm,  which,  how^ever,  continues  to  grow,  so  that  the  cyst  as 
a  whole  increases  in  size.  The  pansporoblasts  at  first  are  uninucleate 
(fig.  112,  a),  and  then  undergo  nuclear  multiplication.  This  is  w^ell  seen 
in  the  intermediate  zone  of  parasites,  where  the  pansporoblasts  show 
first  one,  then  two,  then  four  or  more  spores  (fig.  112,  b),  while  in  the 
oldest  centrally  placed  pansporoblasts,  about  a  dozen  or  sixteen 
closely  packed  spores  (fig.  112,  c),  can  be  seen.  The  spore  is  small 
and  rounded,  and  its  nucleus  is  clear  and  distinct.  The  fully  formed 
pansporoblast  or  spore  morula  becomes  surrounded  by  a  membrane. 

Certain    of  the    cysts  have  been  found  in  a  ruptured  condition,. 

'   QuarL  Journ.  Microsc.  Sci.y  xlix,  p.  521. 


RHINOSPORIDIUM    KINEALYI 


97 


whereby  the  spores  have  been  hberated  into  the  surrounding  tissue. 
It  is  ahiiost  certain  that  the  spores  serve  for  the  auto-infection  of  the 
host,  for  though  the  tumours  of  Rhinosporidium  seemed  to  have 
been  removed  entirely,  it  has  been  found  that  they  recur,  some 
minute  fragment  of  the  parasite  having  probably  been  left  behind. 
The  method  whereby  the  parasite  reaches  new  hosts  has  not  yet  been 
determined,  and  it  would  be  of  niterest  if  its  life-history  could  be 
more  fully  investigated. 


Fig.  112.  —  Rhinosporidium  kinealyi.  Protion  of  ripe 
cyst  containing  pansporoblasts  of  various  ages.  x  480. 
(After  Minchin  and  Fantham.) 

The  Asiatic  specimens  of  R.  kinealyi  were  first  described  in  detail 
by  Minchin  and  Fantham  (1905)  from  material  briefly  reported  to  the 
Laryngological  Society  of  London  in  1903,  by  O'Kinealy.  Material 
obtained  by  Dr.  Nair,  of  Madras,  was  described  by  Beattie^  in  1906. 
This  material  came  from  Cochin.  Castellani  and  Chalmers  have 
found  similar  polypi  in  Ceylon. 

Wright^  has  described  the  parasite  from  Memphis,  Tennessee. 
Seeber^  in  1896  described  nasal  polypi  in  Buenos  Ayres,  and  in  1900 
Wernicke  named  the  parasite  therein  Cocci diiun  seeberi.  Seeber's 
parasite  is  a  Rhinosporidium,  R.  seeberi,  and  may  ultimately  be  found 
to  be  the  same  as  R.  kinealyi,  Ingram*  reports  Rhinosporidium  cysts, 
with  pores  in  the  cyst  walls,  in  conjunctival  polypus  and  in  papilloma 
of  the  penis  in  India.  Zschokke  has  reported  the  presence  of  Rhinos- 
poridiiun  in  horses  in  South  Africa. 


'^Journ.  Pathol,  and  Bacterial.,  xi,  p.  270;  and  ^r?V.  Med.  Journ.,  Nov.  16,  1907,  p.  1402. 

''■  Nezu  York  Med.  Journ.,  December  21,  1907,  p.  1149. 

^  La  Ciencia  Medica  (Buenos  Ayres),  1912.  '  Lancet,  September  3,  1910,  p.  726. 


198  THE    ANIMAL    PARASITES    OF    MAN 

Class  IV.     INFUSORIA,  Ledermuller,  1763. 

The  Infusoria  (or  Heterokaryota,  Hickson,  or  Ciliophora,  Doflein) 
include  the  Ciliata  and  the  Suctoria.  A  few  authorities,  including 
Braun,  raise  the  Suctoria  (or  Acinetaria)  to  separate  rank  as  a  class, 
but   this  is  not  widely   followed. 

The  body  of  the  Ciliata  usually  is  bilaterally  symmetrical  and  is. 
enveloped  in  a  cuticle  which  has  numerous  openings  for  the 
protrusion  of  the  cilia.  Most  kinds  have  a  fixed  shape,  whilst 
changes  in  the  form  of  others  are  brought  about  by  the  contractions 
of  the  body  substance.  The  latter  exhibits  hyaline  ectoplasm,  in 
which  myonemes,  and  occasionally  also  trichocysts  (minute  spindle- 
shaped  bodies)  appear,  and  granular  endoplasm  which  may  contain 
numerous  vacuoles.  The  cilia,  on  whose  various  arrangements 
the  classification  is  based,  are  always  processes  of  the  ectoplasm. 
Their  form  varies ;  they  may  be  hair-like,  or  more  rarely  thorn-like, 
spur-like,  or  hook-shaped ;  undulatory  membranes  also  may  occur, 
which  are  probably  composed  of  fused  cilia. 

With  the  exception  of  some  of  the  parasitic  species,  an  oral  cavity, 
peristome  or  cytostome,  is  always  present.  It  is  frequently  beset 
with  cilia  or  provided  with  undulatory  membranes,  which  help  to 
waft  the  food  inwards;  sometimes  there  is  an  anal  aperture  (cytopyge) 
generally  placed  at  the  opposite  pole  of  the  organism.  A  cyto- 
pharynx  fringed  with  cilia  or  sometimes  with  a  specialized  supporting 
apparatus  is  connected  with  the  peristome.  Vacuoles  form  round 
the  ingested  food,  and  in  many  species  a  constant  rotation  goes  on 
in  the  endoplasm.  Often  one,  and  sometimes  two  contractile  vacuoles 
are  present,  the  frequency  of  the  pulsations  of  which  depends  on  the 
surrounding  temperature.  Sometimes  special  conducting  channels 
lead  to  the  vacuoles,  or  there  are  outlet  channels  leading  to  the  exterior. 

There  is  in  almost  every  case  a  large  nucleus  (macronucleus), 
and  lying  close  up  to  it  a  small  nucleus  (micronucleus).  The  form 
of  the  large  nucleus  varies  according  to  the  species.  Numerous 
nuclei  are  not  very  common,  but  these  occur  in  Opalina,  which  lives 
in  the  hind-gut  of  amphibia,  and  is  also  distinguished  by  the  absence 
of  an  oral  aperture. 

Reproduction  is  effected  by  binary  fission  ;  less  commonly,  after 
encystment,  by  multiple  division,  or  by  budding.  The  divisions  can 
be  repeated  many  times,  but  finally  cease,  and  then  the  conjugation 
of  two  specimens  brings  about  a  regeneration,  particularly  of  the 
nuclei.  Numerous  examinations  (Biitschli,  Hertwig,  Maupas,  Calkins) 
have  demonstrated  that  after  tw^o  individuals  have  associated  by 
homologous  parts  of  the  body,  the  micronucleus  separates  from 
the  macronucleus,  becomes  larger  and  divides  twice  by  mitosis,  so 


INFUSORIA 


199 


that  four  microniiclei  are  present  in  each  one  of  the  two  individuals 
forming  the  couple.  Three  of  these  nuclei  perish  and  become 
absorbed,  the  fourth  gradually  passes  to  the  portion  of  protoplasm 
connecting  the  two  conjugants,  which  has  originated  by  absorption 
of  the  cuticle  at  the  point  of  contact  of  the  conjugants.  After  a 
further  division  one  micronucleus  of  each  conjugant  passes  over 
into  the  other  conjugant,  and  fusion  ensues  between  the  two  micro- 
nuclei  of  each  individual.  Complicated  changes  and  divisions  may 
occur,  but  only  the  main  principles  can  be  noted  here.  A  new  nuclear 
body  is  thus  formed  in  each  conjugant,  and  soon  divides  into  two. 
Of  the  segments  thus  produced  one  becomes  a  micronucleus,  and 
one  or  several  of  the  others,  as  the  case  may  be,  form  or  amalgamate 
into  a  new  macronucleus,  the  old  macronucleus  usually  perishing  or 
becoming  absorbed  during  the  conjugation.  Usually,  sooner  or  later, 
the  two  conjugants  separate,  or  may  have  separated  already,  and  again 
multiply  independently  by  fission  until  a  series  of  divisions  by  simple 
fission  is  again  followed  by  conjugation.  The  theoretical  signifi 
cance  of  conjugation  cannot  be  dealt  with  fully  here.  It  may  be 
remarked,  however,  that  the  macronucleus  plays  no  part  in  it,  but 
governs  entirely  the  metabolism  of  the  Infusorian,  whereas  the 
micronucleus  is  essentially  a  generative  nucleus  from  which  macro- 
and  micro-nuclei  are  again  and  again  produced. 

Encystment  amongst  the  Infusoria  is  very  general,  and  is  essentially 
a  means  of  protection  when  the  surrounding  medium  dries  up. 
Doubtless  these  cysts  are  frequently  carried  long  distances  by  the  wind, 
which  explains  the  wide  geographical  distribution  of  most  species. 
Also,  multiplication  often  takes  place  in  the  encysted  condition. 

Some  Infusoria  live  a  free  life,  others  are  sedentary;  the  latter 
form  colonies  in  fresh  as  well  as  in  salt  water.  Numerous  species 
are  parasites  of  various  lower  and  higher  animals,^  and  a  few  also  are 
parasitic  in  man. 

The  Prague  zoologist,  v.  Stein,  introduced  a  classification  of  the 
Infusoria  that  has  been  almost  universally  adopted.  It  is  founded 
on  the  dififerent  position  of  the  cilia  on  the  body.  Though,  no 
doubt,  artificial,  it  is  a  convenient  system.  Biitschli  has  compiled  a 
better  one.      But  for  our  purpose  Stein's  system  is  sufficient : — 

Order  i.     Holotricha^  Infusoria  with  cilia  that  are  evenly  distributed  over  the 
entire  body. 


'  It  may  be  stated  that  numerous  peculiarly  shaped  species  live  in  the  stomach  of  rumi- 
nants, others  in  the  colon  of  horses.  Several  species  are  found  in  the  rectum  of  frogs  and 
toads ;  others,  again,  on  the  surface  of  the  bodies  of  fishes  ;  and  various  other  species  exist 
in  and  on  the  bodies  of  invertebrate  animals. 

-  Bronn's  CI.  u.  Ordn.  d.   Thierr.,  i.  Protozoa,  Part  3,  Infusoria. 


200 


THE   ANIMAL   PARASITES   OF   MAN 


Order  2.     Heterotricha,  ciliated  all  over  like  the  Holotricha,  but  having  stouter 

cilia  about  the  peristome. 
Order  3.     Hypotricha^  only  ciliated  on  the  ventral  surface. 
Order  4.     Periiricha,  with  only  a  ring  of  spiral  cilia,  mostly  sedentary. 

The  Infusoria  observed  in  man  belong  to  the  order  HeterotricJui, 
with  few  exceptions. 

Genus.  Balantidium,  Claparede  et  Lachmann. 
Heterotrichous  Infusoria  of  oval  or  bag-like  form  and  almost  circular  on  trans- 
verse section  ;  the  anterior  extremity  narrowed,  the  posterior  end  broad  and 
rounded  off,  or  also  narrowed  ;  the  peristome  starting  at  the  anterior  end  is 
broadest  there  and  becomes  narrower  as  it  gradually  obliquely  approaches  towards 
the  posterior  extremity.  There  are  coarse  cilia  along  the  entire  left  border  and  the 
anterior  part  of  the  right  border.  Longitudinal  striation  is  distinct  and  regular. 
There  are  two  contractile  vacuoles  on  the  right,  and  occasionally  also  two  or  more  to 
the  left.  The  anus  (cytopyge)  is  terminal.  The  macronucleus  is  usually  horse-shoe 
or  kidney-shaped,  sometimes  oval ;  the  micronucleus  contiguous.  Reproduction  is 
by  binary  fission  and  conjugation,  and  encystment  occurs.  The  cysts  are  spherical 
or  oval.  These  ciliates  are  parasitic  in  the  large  intestine  of  human  beings  and 
pigs,  in  Amphibia,  and  in  the  body  cavity  of  polychaste  Annelida. 

Balantidium  coll,  Mahiisten,  1857. 

Syn. :    ParamcEcium  coli^  Malmsten,  1857. 

The  body  is  oval,  60  ^L  to  100  /a  in  length  (up  to  200  yu  according 
to  Janowski),  and  50  //,  to  70  /a  in  breadth.  The  peristome  is  funnel- 
shaped   or   contracted,  the  anterior   end  being    then    broadened  or 


Fig.  113. — Balati- 
Hdiuvi  coli.  a,  nu- 
cleus ;  d,  vacuole  ; 
c,  peristome  ;  d,  bolus 
of  food.  (After 
Leuckart.) 


Fig.  114. — Balantidium  coli,  free  and  encysted; 
a,  anus  or  cytopyge ;  n,  macronucleus  :  />,  bolus  of 
food.     (After  Casagrandi  and  Barbagallo.) 


pointed  according  to  the  degree  of  contraction  (figs.  113,  114).  The 
ecto-  and  endo-plasm  are  distinct,  the  latter  is  granular,  containing 
drops  of  fat  and  mucus,  granules  of  starch,  bacteria,  and  occasionally 


BALANTIUIUM    COLI  20I 

also  red  and  white  blood  corpuscles.  There  are  usually  two  con- 
tractile vacuoles,  seldom  more.  The  anus  (cytopyge)  opens  at  the 
posterior  extremity.  The  macronucleus  is  bean-  or  kidney-shaped, 
rarely  oval ;  the  micronucleus  is  spherical. 

Balaiitidium  coll  lives  in  the  large  intestine  of  man,  in  the  rectum  of 
the  domestic  pig,  and  has  been  found  in  monkeys.  It  propagates  by 
transverse  division,  but  conjugation  and  encystment  are  known  to 
take  place.^  Transmission  to  other  hosts  is  eft'ected  by  the  cysts  of 
the  parasite  (fig.  114). 

Balaiitidinjn  coll,  first  seen  by  Leeuwenhoek,  was  described 
by  Malmsten  in  1857  in  a  man  aged  35  years,  who  had  two 
years  previously  suffered  from  cholera,  and  since  then  had  been 
subject  to  diarrhoea.  The  examination  showed  an  ulcer  in  the  rectum 
above  the  mid  sphincter  ani,  in  the  sanguineous  purulent  secretion  of 
which  numerous  Balantidia  were  swimming  about.  Although  the 
r  deer  was  made  to  heal,  the  diarrhoea  did  not  cease  and  the  stools 
contained  numerous  Balantidia,  the  number  of  which  could  only  be 
decreased  by  extensive  enemas  of  hydrochloric  acid. 

The  second  case  related  to  a  woman  who  was  suffering  from 
severe  colitis,  and  who  died  ten  days  after  admission.  The  mal- 
odorous, watery  evacuations  contained  innumerable  Balantidia,  in 
addition  to  pus,  and  at  the  autopsy  the  anterior  portion  of  the  large 
intestine  was  found  to  be  infested  with  them. 

Subsequently  this  parasite  has  often  been  observed  in  human 
beings,  and  various  cases  have  been  recorded.  These  occurred  in 
Russia,  Scandinavia,  Finland,  Cochin  China,  Italy,  Germany,  Serbia, 
Sunda  Islands,  Philippine  Islands,  China,  and  in  other  parts  of  Asia 
and  in  America.  Other  cases  were  reported  by  Askanazy,  Ehrnroth, 
Klimenko,  Nagel,  Koslowsky,  Kossler,  Waljeff,  Strong  and  Musgrave, 
Glaessner^  and  others.     Sievers  found  B.  coll  very  common  in  Fmland. 

In  the  majority  of  the  cases  described  by  Sievers  from  Finland, 
and  in  other  cases  from  Central  Europe,  the  patients  suffered  from 
obstinate  intestinal  catarrh,  w^hich  did  not  always  cease  even  after 
the  Balantidia  had  disappeared.  On  the  other  hand,  Balantidia  have 
occasionally  still  been  found  to  persist,  though  in  small  numbers,  after 
the    catarrh    has   been    cured.     Some   authors,    nevertheless,    do    not 

•  According  to  Gourvitch  ("  Bal.  coli.  Darmk.  d.  Menschen,"  /vnss.  Arch.  f.  Path.^  klin. 
med.  u.  Bad.,  Petrograd,  1896),  the  conjugated  Balantidia  are  supposed  to  fuse  with  each 
other  and  form  oval  cysts  two  or  three  times  the  size  of  the  free  organisms,  and  to  divide  into 
numerous  globules  within  the  cystic  membrane  ;  the  process,  however,  has  hitherto  not  been 
confirmed.  The  supposed  Balantidium  cysts  appeared  in  two  patients  who  were  simultaneously 
suffering  from  Dibothriocephahis  latus,  after  the  administration  of  anthelminthics.  It  therefore 
seems,  according  to  the  description,  that  in  reality  these  forms  were  actually  abnormally  large, 
possibly  swollen,  young  eggs  of  the  tape-worm  mentioned. 

•  Centralbl.f.  Rakt.,  Orig.,  xlvii,  p.  351. 


202  THE    ANIMAL    PARASITES    OF   MAN 

regard  Balantidia  as  the  primary  cause  of  the  various  diseases  of  the 
large  intestine,  which  often  commence  with  the  development  of 
ulcers,  but  they  consider  that  they  may  aggravate  these  diseases  and 
render  them  obstinate.  According  to  Solowjew,  Askanazy,  Klimenko 
and  Strong  and  Musgrave,  however,  the  parasites  penetrate  the  intes- 
tinal wall,  and  give  rise  to  ulcerations  which  may  extend  deeply  into 
the  submucosa,  and  even  be  found  in  the  blood  and  lymphatic  vessels 
of  the  intestinal  wall.  According  to  Stokvis,  B.  coli  occurs  also  in 
the  lung ;  at  all  events  this  author  states  that  he  found  one  living  and 
several  dead  param^cia  (?)  in  the  sputum  of  a  soldier,  returned  from 
the  Sunda  Islands,  who  was  suffering  from  a  pulmonary  abscess. 
Sievers  has  shown  that  B.  coli  might  occur  in  persons  not  suffering 
from  intestinal  complaints,  but  E.  L.  Walker  ^  (19^3)  states  that  every 
person  parasitised  with  B..  coli  is  liable  sooner  or  later  to  develop 
balantidian  dysentery. 

Since  Leuckart  confirmed  the  frequent  presence  of  B.  coli  in  the 
rectum  of  pigs,  and  corresponding  observations  were  made  in  other 
countries,  the  pig  is  universally  considered  to  be  the  means  of  the 
transmission  of  Balantidium  to  man.  The  encysted  stages  only  serve 
for  transmission,  because,  according  to  all  observations,  the  free 
parasites  have  a  very  small  power  of  resistance.  They  perish  when 
the  faeces  have  become  cool ;  they  cannot  live  in  ordinary,  slimy,  or 
salt  water.  As  they  are  killed  by  acids  even  when  much  diluted,  they 
cannot  pass  through  the  normal  stomach  alive  except  under  the  most 
unusual  circumstances.  The  pigs,  in  whose  intestines  the  Balantidium 
appears  to  cause  little  or  no  disturbance,  evacuate  numerous  encysted 
Balantidia  with  the  faeces,  and  their  occasional  transference  to  man 
brings  about  their  colonization  there,  but  perhaps  only  when  a  disease 
of  the  colon  already  exists. 

Experimental  transmission  of  the  free  parasites  to  animals  (per  os 
or  per  anum)  yielded  negative  results,  even  in  the  case  of  pigs. 
Casagrandi  and  Barbagallo  (1896),  however,  had  positive,  as  well  as 
negative,  results.  They  employed  healthy  young  cats,  or  cats  in  which 
catarrhal  entero-colitis  had  been  artificially  induced  (which  in  other 
experiments  is  apt  to  cause  the  death  of  the  animals  experimented  upon 
in  about  six  or  seven  days),  or  finally  cats  that  had  dilatation  of  the 
rectum  with  alkaline  reaction  of  the  faeces.  An  attempt  to  infect  three 
healthy  cats  by  injecting  human  faeces  containing  Balantidium  into  the 
rectum  proved  negative,  in  so  far  as  the  faeces  of  the  experimental 
animals  had  an  acid  reaction  and  contained  no  Balantidia,  but  at  the 
autopsy  performed  eight  days  after  infection  a  few^  encysted  parasites 
were  found  in  the  mucus  of   the  ileum.     In  the  case  of   four  cats 

^  Philip.  JL  Sc,  Sec.  B,  viii,  p.  333. 


BALANTIDIUM    COLI 


203 


suffering  from  entero-colitis,  into  which  human  faeces  containing 
Balantidia  were  introduced  per  os,  Balantidium  cysts  were  found  in 
the  faeces  three  days  after  the  last  ingestion.  Great  numbers,  more- 
over, w^ere  found  in  the  caecum  and  the  posterior  part  of  the  small 
intestine  at  the  autopsy  of  the  animals,  which  died  about  eight  days 
after  the  commencement  of  the  experiments.  Actual  colonization, 
therefore,  was  not  effected  in  either  series  of  experiments.  Free  or 
encysted  Balantidia  of  pigs  were  used  for  further  experiments.  The 
experiments  proved  negative  when  faeces  containing  cysts  were  injected 
into  the  rectum  of  healthy  cats  (three  experiments),  or  cats  (two) 
suffering  from  spontaneous  intestinal  catarrh,  or  when  such  material 
was  introduced  per  os  into  three  healthy  cats.  In  the  case  of  two 
cats  with  intestinal  catarrh  artificially  produced,  a  small  number  of  the 
active  Balantidia  injected  into  the  rectum  remained  alive.  Larger 
quantities  of  faeces  containing  encysted  Balantidia  were  introduced 
into  two  other  cats  affected  with  the  same  complaint.  These,  certainly, 
did  not  appear  in  the  faeces,  but  small  numbers,  free  and  alive,  were 
found  in  the  caecum.  Similarly,  encysted  Balantidia  were  introduced 
into  two  cats  with  dilated  rectum,  and  whose  faeces  had  an  alkaline 
reaction,  in  these  cases  no  parasites  appeared  in  the  faeces,  but  three 
and  five  days  later,  when  the  two  animals  were  examined,  a  very  small 
number  were  discovered  free  in  the  large  intestine.  Klimenko  did 
not  succeed  in  infection  experiments  with  B.  coll  on  young  dogs, 
whose  intestines  had  been  artificially  affected  by  disease. 

More  recent  experiments  by  Brumpt  have  shown  that  young 
sucking  pigs  can  be  infected  with  Balantidium  from  infected  monkeys 
{Macacns  cynoniolgiis)  and  suffer  heavily  from  the  same,  whereas  the 
Balantidium  of  the  pig  is  rarely  harmful  to  its  host.  This  and  previous 
experiments  may  be  thought  to  suggest  that  there  are  perhaps 
several  pathogenic  species,  and  also  that  harmless  strains  of  Balan- 
tidium may  occur.  At  the  same  time,  it  must  be  remembered  that 
a  large  proportion  of  the  cases  recorded  of  Balantidian  colitis  occur 
among  swineherds  and  butchers,  that  is,  among  people  in  frequent 
contact  with  pigs.  Morphologically,  there  are  practically  no  differ- 
ences between  the  Balantidia  found  in  man,  monkeys  and  pigs,  and 
it  is  probable  that  one  species  only,  under  slightly  different  environ- 
mental conditions,  may  be  responsible  for  the  colitis  observed.  In 
any  case,  efficient  prophylactic  measures  should  be  taken  against 
balantidiasis  in  countries  where  it  may  occur,  by  confining  the  pigs 
and  not  allowing  them  to  run  in  yards  and  dwellings. 

E.  L.  Walker  (19 13)  has  given  a  good  summary  of  work  on 
balantidiasis.  His  own  researches  in  the  Philippines  showed  that 
monkeys  could  be  infected  by  Balantidia  both  from  pigs  and  men. 
Parasites  may  appear  in  the  stools  only  at  infrequent  intervals.     He 


204 


THE    ANIMAL   PARASITES   OF   MAN 


believes    that   the    cihates  are    the    prhiiary   etiologic    factor    in    the 
symptoms  and  lesions  of  balantidian  dysentery. 

Behrenroth  (1913)  has  given  an  interesting  account  of  Balantidium 
coli  and  its  pathogenic  significance. 


Balantidium    minutum,  Schaudinn,  1899. 

The  body  is  of  oval  form,  with  tlie  anterior  extremity  pointed,  and 
posterior  extremity  broad  and  rounded  (fig.  115).  The  length  is  20  fi 
to  32  /jL,  and  the  breadth  is  14  /jl  to  20  /x.  The  peristome,  which  is 
fissure-like,  extends  to  the  centre  of  the  body  (fig.  115).  The  right 
lateral  border  of  the  peristome  is  beset  with  cilia  the  same  length  as 
those  of  the  body,  the  left  side  terminates  in  a  thin  hyaline  membrane 

that  extends  towards  the  back,  and  can 
pass  over  to  the  right  side.  A  row  of 
longer  and  stronger  cilia  (cirri)  are  on  the 
left  border  of  the  peristome.  The  cuticle 
is  refractile,  the  ectoplasm  hyaline  and 
the  endoplasm  granular,  with  numerous 
food  vacuoles. 

A  single  contractile  vacuole  lies  dor- 
sally  and  to  one  side  at  the  posterior 
extremity.  The  macronucleus,  which  is 
always  spherical,  is  central  and  is  6  /x  to 
7/x  in  diameter.  The  micronucleus,  close 
in  front  of  it,  only  measures  i  /a  (fig.  115). 
The  cysts  are  oval. 

These  parasites  were  found  in  num- 
bers in  the  evacuations  of  a  man  aged  30, 
who  was  born  in  Germany  and  had  repeatedly  travelled  between 
Hamburg  and  North  America,  where  he  made  long  stays.  Tlie  patient 
came  to  the  Charite  in  Berlin  to  seek  advice  for  constipation  alter- 
nating WTth  diarrhoea  accompanied  by  abdommal  pam. 

A  second  case  (the  parasite  of  which  w^as  described  as  Colpoda 
<:iiciilliis  by  Schulz)  was  observed  in  a  patient  in  the  same  institution. 
As,  in  both  cases,  the  parasites  only  appeared  during  the  diarrhoea, 
and  disappeared  as  soon  as  the  faeces  had  assumed  a  normal  consis- 
tency, or  could  only  be  demonstrated  in  a  few  encysted  specimens, 
it  may  be  assumed  that  the  small  intestine  or  the  duodenum  is 
their  habitat. 


Fig.  115. — Balantidium  minu- 
itim.  P,  peristome;  N,  nucleus; 
J/,  micronucleus ;  V,  contractile 
vacuole.  Food  vacuoles  are  repre- 
sented in  the  endoplasm.  (After 
Schaudinn.) 


Genus.     Nyctotherus,  Leidy,  1849. 

Flat,   heterotrichous    Infusoria,   kidney-  or   bean-shaped.     The   peristome    com- 
mences at  the  anterior  pole  of  the  body  and  extends  along  the  concave  side  to  the 


NYCTOTHERUS   FAB  A 


^05 


middle,  where  the  oral  aperture  is  situated.  The  cytopharynx  is  oblique  and  is  more 
or  less  curved.  The  cytopyge  is  at  the  posterior  extremity,  where  a  single  contractile 
vacuole  is  also  situated.  The  macronucleus  is  almost  in  the  centre  of  the  parasite. 
The  members  of  this  genus  live  parasitically  in  the  intestine  of  amphibia,  insects 
and  myriapods,  and  at  least  one  species  is  found  in  man. 


Nyctotherus  faba,  Schaudinn,  1899. 

The  body  is  bean-shaped,  and  a  Httle  flattened  dorso-venlrally. 
It  is  26  //,  to  28  //,  ^ong,  16  yL6  to  18  yLt  bi'oad,  and  10  //,  to  12  /a  thick 
(fig.  116).  The  peristome  is  on  the  right 
border  and  extends  to  the  middle ;  at 
the  left  there  are  large  adoral  cilia,  the 
cilia  on  the  right  border  not  being  larger 
than  those  on  the  body.  The  cyto- 
pharynx is  short,  slightly  curved  and 
turned  backwards.  The  contractile  vac- 
uole is  large,  spherical,  situated  at  the 
posterior  extremity,  and  its  contents  are 
voided  through  the  anus  at  its  left.  The 
macronucleus  is  in  the  centre  of  the  body; 
it  is  globular  (6  /x  to  7  //,  in  size),  and 
contains  four  or  five  chromatin   masses.        ¥ig.  116.— JVyao/Aerus fada.  r, 

rr^.  .  ,  ,.  1  ,       .,  ,    .         peristome;  A^.  nucleus ;  ill/,  micro- 

1  he   micronucleus  lies  close    to    it,  and    is       nucleus;     r,    contractile    vacuole. 

spherical  or  somewhat  elongate  measuring     (After  Schaudinn.) 
I  /A  to  1*5  w,  (fig.  116).     The  cysts  are  oval. 

This  species  has  hitherto  only  been  seen  once  in  the  same  patient 
in  whom  Balantiduim  niinutum  was  discovered. 


Nyctotherus   giganteus,  P.  Krause,  1906. 

Under  the  name  Balaiifidiumgigaiiteiun  n.  sp.,  P.  Krause  described 
an  Infusorian  which  was  repeatedly  observed  with  Trichoiuonas  iutes- 
fiiialis  in  the  alkaline  evacuations  of  a  typhoid  patient  in  Breslau. 
The  body  is  ovoid,  narrower  and  rounded  anteriorly  and  broader 
and  stunted  posteriorly.  The  peristome  lies  to  one  side  ;  the  macro- 
nucleus  is  bean-shaped,  the  micronucleus  small  and  globular ;  one 
or  two  vacuoles  are  present.  The  anus  is  at  the  farther  end.  The 
organism  is  go  /jl  to  400  /^  long,  60  fi  to  150  fju  broad  (fig.  117).  After  a 
prolonged  stay  outside  the  body,  it  becomes  rounded  and  encystment 
occurs.  In  the  thermostat  the  Infusoria  remain  alive  at  37°  C.  for  five 
weeks. 

The  species,  however,  hardly  belongs  to  Balautidiiuii,  but  to  all 
appearances  is  a  Nyctoihents  and  is  distinguished  from  N.  faha  by  the 
difference  in  size. 


>o6 


THE    ANIMr^L    PAI^ASITES   OF    MAN 


[Nyctotherus]  afrlcanus,  Castellani,   1905. 

In  the  fa3ces  of  a  native  of  Uganda  who  suffered  froni  sleeping 
sickness  and  diarrhoea  and  had  in  his  intestine  Ascaris  liujibricoidcs, 
Trichoceplialus  trichiiinis  and  Ancylostoiiui  diiodenale,  Castellani  found 
a  curiously  shaped  Infusorian,  40  /x  to  50  /x  long,  and  35  //,  to  40  />t 
broad,  with  spherical  macro-  and  micronucleus  and  a  contractile 
vacuole  (tig.  118).  He  included  the  organism  in  the  genus  Nyctotherus, 
perhaps  wrongly,  or  the  parasite  may  have  been  deformed.  After  the 
patient's  death  the  same  parasite  was  found  in  the  intestine  and 
especially  in  the  caecum. 


Fig.    WJ.— Nycto- 
therus giganleiis. 
(Afier  Krause.) 


Fig.  i\%.— Nyctotherus 
africamis.  (After  Castel- 
lani.) 


G.  Lindner,  in  Cassel,  studied  certain  peritrichal  Infusoria  (stalkless  Vorticella), 
and  connected  them,  probably  incorrectly,  with  the  most  varied  diseases  of  man  and 
domestic  animals,  even  with  Sarcosporidia  of  pigs.  It  may  be  mentioned  that 
according  to  a  communication  by  letter  from  Schaudinn,  Vorticella  may  be  found 
in  freshly  evacuated  faeces,  but  always  only  after  the  administration  of  a  water 
enema.  In  spile  of  this,  several  other  investigators  mention  Vorticellas  as 
intestinal  parasites  of  man. 

The  Chilodon  dentafus  (Ehrenberg)  recorded  in  1903  by  J.  Guiart  as  a  parasite 
of  man,  which  may  be  found  in  all  infusions,  can  hardly  have  lived  in  the  man 
from  whose  faeces  it  was  cultivated,  but  may  represent  a  chance  admixture  both  in 
the  faeces  and  the  cultivations.  C.  uncifiahis  was  also  found  as  a  chance  parasite 
of  man  by  Manson  and  Sambon.  According  to  Doflein^  (1911)  certain  Chilodon- 
like  organisms  have  been  found  by  Selenew  in  prostate  secretions  in  gonorrhoea. 
Other  species  of  the  genus  Chilodon  are  known,  but  only  as  ectoparasites  {e.g., 
Chilodo?i  cypriiii,  Morofif,  1902,  from  the  skin  and  gills  of  diseased  carp). 

A  number  of  other  parasitic  Ciliates  are  known,  among  which  Ichthyophthirius 
jiiultifiliis,  destructive  to  fish,  is  important.  It  lives  in  the  skin  and  the  layers 
immediately  below  it,  forming  small  whitish  pustules  which  may  become  confluent. 


'   Lehrhuch  der  Frotozoeukiinde,  3rd  ed,,  p.  963. 


CHLAMYDOZOA  20/ 

The  pustules  are  most  common  on  the  head  and  fins,  but  occur  also  on  the  eyes  and 
gills  of  the  host.  The  young  parasite,  which  is  one  of  many  formed  in  a  cyst,  is  very 
small.  At  first  it  is  free  swimming,  but  soon  attaches  itself  to  the  skin  of  a  fish.  It 
bores  inwards  and  becomes  surrounded  by  the  irritated  skin.  There  it  attains  a 
relatively  large  size,  being  500 /a  to  750 /^  and  occasionally  more  in  diameter.  The 
body  has  a  rounded  terminal  mouth,  short  cytopharynx  and  a  number  of  minute 
contractile  vacuoles.  The  macronucleus  is  large  and  horseshoe-shaped  ;  the  small 
micronucleus  is  only  seen  in  the  very  young  animal.  When  full  grown,  the  organism 
encysts  and  forces  its  way  to  the  surface  and  bursts  through,  leaving  a  small,  gaping 
wound  behind.  The  cyst  sinks  to  the  bottom  of  the  water,  nuclear  multiplication 
occurs  and  a  number  of  young  parasites  are  produced,  which  leave  the  cyst  and 
either  attack  new  hosts  or  else  perish. 

Opalina  ranai'um,  parasitic  in  the  rectum  and  urinary  bladder  of  frogs  and  toads, 
shows  great  degradation  and  simplification  due  to  parasitism,  possessing  no  separate 
micronuclei,  no  cytostome,  cytopharynx  or  cytopyge.  It  has  many  macronuclei,  and 
is  a  large  parasite.  During  summer  and  autumn  nuclear  multiplication  followed  by 
division  of  the  body  occurs,  the  process  being  repeated  after  the  daughter  forms 
have  grown  to  the  size  of  their  parent.  In  spring,  the  Opalina  divide  rapidly,  but 
do  not  grow  much  before  dividing  again.  Finally,  tiny  forms,  containing  three  to 
six  nuclei,  encyst  and  pass  from  the  host  with  the  fteces.  As  these  latter  are  greedily 
devoured  by  tadpoles,  the  Opalina  gain  new  hosts  in  which  they  develop. 


THE  CHLAMYDOZOA. 

The  name  Chlamydozoa  was  proposed  by  Prowazek  in  1907  for  a 
number  of  minute,  problematic  organisms  (fig.  119)  believed  to  be 
the  causal  agents  of  certain  diseases  in  man  and  animals,  such  as 
vaccinia  and  variola,  trachoma,  inclusion  blenorrhoea  in  infants, 
molluscum  contagiosum,  and  bird  epithelioma  contagiosum.  Other 
diseases  possibly  due  to  Chlamydozoa  ^  are  hydrophobia,  measles, 
scarlet  fever,  foot-and-mouth  disease,  the  "  Gelbsucht"  disease  of  silk- 
worms, and  perhaps  even  typhus  (Prowazek,  191 3).  The  subject  is 
difficult  and  controversial  and  can  only  be  briefly  discussed  here. 
It  is  known  that  the  viruses  in  all  these  diseases  can  pass  through 
ordinary  bacterial  filters,  that  is,  they  belong  to  the  group  of  ''  filterable 
viruses."  At  such  periods  the  organisms  are  extracellular  or  free.  It 
is  also  known  that  in  many  of  these  cases  the  virus  produces  definite 
and  characteristic  reaction-products  or  cell-inclusions  in  the  infected 
cells,  during  the  intracellular  phase  of  the  life-history  of  the  organism. 
As  the  organisms  to  be  considered  are  problematic,  it  will  be  con- 
venient to  summarize  their  history  : — 

(i)  Cell-inclusions,  usually  named  after  their  discoverers,  have 
been  found  in  certain  diseases,  thus  :   In  vaccinia  Guarnieri's  bodies, 

'  For  a  detailed  account  of  the  Chlamydozoa  see   Prowazek's  Handbtich  der  Pathogenen 
Frotozoen,  Bd.  i  (191 1- 12).     Leipzig,  J.  A.  Barth. 


208  THE    ANIMAL    PARASITES    OF   MAN 

in  scarlet  fever  Mallory's  bodies,  in  hydrophobia   Negri's  bodies,  in 
trachoma  Prowazek's  bodies  occur. 

(2)  At  first  these  characteristic  cell-inclusions  were  considered  to 
be  actual  parasitic  organisms  causing  the  diseases  in  question.  The 
bodies  received  zoological  names  and  attempts  were  made  to  work 
out  their  supposed  development  cycles.  The  supposed  parasites  of 
vaccinia  and  variola  were  referred  to  a  so-called  genus  Cytorydes, 
those  of  hydrophobia  to  Neuroryctes,  of  scarlet  fever  to  Cyclasierhim, 
while  those  of  molluscum  contagiosum  were  referred  to  the  Coccidia. 
Calkins  in  1904  studied  in  detail  the  cell-inclusions  of  vaccinia  and 
small-pox,  calling  them  Cytorydes  varlolce,  Guarnieri.  Calkins  con- 
sidered that  in  the  stratified  cells  of  the  epidermis  they  passed  through 
two  cycles,  the  one  cytoplasmic,  the  other  intranuclear.  The  first  is 
the  vaccinia  cycle,  the  second  the  pathogenic  (intranuclear)  variola 
cycle.     It   is   hardly   necessary   to    follow   all    Calkins'    stages    here. 

Negri  (1909)  described  a  cycle  for  Nenrorydes  hydrophobia.  Calkins 
refers  both  Cytorydes  variotcc  and  Neuroryctes  hydrophohice  to  the 
Rhizopoda. 

Siegel  (1905)  described  quite  different  organisms  under  the  name 
Cvtorhydes.  He  listed  several  species  :  C.  vacdnice  of  vaccinia  and 
small-pox,  C.  scarlatlnce  of  scarlet  fever,  C.  luis  of  syphilis  (this  is 
probably  the  granule  stage  of  Treponema  pallidiun),  and  C.  aphtharuju 
of  foot-and-mouth  disease. 

(3)  The  afore-mentioned  views  were  criticized,  and  the  bodies 
were  not  considered  to  be  living  organisms  but  merely  reaction 
products  or  cell-inclusions  due  to  the  effects  of  the  virus  on  the  host 
cells.  Thus  Guarnieri's  bodies  were  stated  to  consist  of  extruded 
nucleolar  or  plastin  material,  having  no  developmental  cycle.  It  was 
further  asserted  that  infection  could  be  produced  by  lymph  in  which 
Guarnieri's  bodies  had  been  destroyed.  Similar  assertions  have  been 
made  regarding  the  Negri  bodies,  and  others.  The  Cytorydes, 
Neurorydes,  etc.,  are  considered,  according  to  these  views,  to  be 
degeneration  products  of  the  nucleus  or  to  be  of  a  mucoid  nature. 

(4)  More  recently  a  positive  belief  has  gained  ground  that  there 
are  true  parasitic  organisms  causing  these  diseases,  and  that  the 
parasites  are  very  minute,  being  termed  Chlamydozoa  by  Prowazek 
and  Strongyloplasmata  by  Lipschlitz. 

The  Chlamydozoa  are  characterized  by  {a)  their  very  minute  size, 
smaller  than  any  bacteria,  so  that  they  can  pass  through  bacterial 
filters ;  (6)  they  pass  through  intracellular  stages,  in  the  cytoplasm 
or  the  nucleus  of  the  host  cell,  producing  therein  the  reaction 
products  or  inclusions  in  the  cell  already  recorded  as  characteristic 
or  diagnostic  of  the  diseases  produced;  (c)  they  pass  through  definite 
developmental   cycles.     Such  a   cycle   consists  essentially  of  growth 


CHLAMYDOZOA 


209 


and  division.  The  mode  of  division  of  the  Chlamydozoa  resembles 
that  of  the  centriole  of  a  cell,  by  the  formation  of  a  dumb-bell-shaped 
figure.  Two  dots  are  observed  connected  by  a  fine  line  or  strand 
which  becomes  drawn  out  and  finally  snaps  across  the  middle. 
Prowazek  and  Aragao  (1909)  working  on  smallpox  in  Rio  de  Janeiro 
found  that  the  chlamydozoal  granules  passed  through  a  Berkefeld 
filter  and  that  the  filtrate  was  virulent.  But  if  an  "ultra-filter"  were 
used,  i.e.,  one  coated  with  agar,  then  the  granules  were  retained  and 
the  filtrate  was  no  longer  virulent.  The  surface  of  the  ultra-filter  was 
found  to  contain  many  granules. 

The  Chlamydozoa  are  parasites  of  epiblastic  tissues  {e.g.y  epidermal 
cells,  nerve  cells,  conjunctival  cells). 

The  life-history  of  a  Chlamydozoon    (fig.    119),  such  as   that   of 


Fig.  119. — Chlamydozoa.  Trachoma  bodies  in  infected 
epithelial  cells  of  the  conjunctiva,  (a)  initial  bodies  (above) 
and  cluster  of  elementary  bodies  (touching  the  nucleus) ; 
{b)  cluster  of  granules  surrounded  by  mantles,  x  2,000 
approx.    (Original.     From  preparation  by  Fantham.) 

vaccinia,  is,  according  to  Prowazek,  Hartmann  and  their  school,  as 
follows  : — 

1.  The  infection  begins  with  elementary  bodies  or  elementary 
corpuscles  which  live  at  first  extracellularly.  An  elementary  body  is 
a  minute  speck  of  chromatin,  apparently  devoid  of  cytoplasm,  which 
can  pass  through  a  bacterial  filter.  It  can  enter  a  host  cell,  but  the 
entry  is  not  a  process  of  phagocytosis. 

2.  Inside  the  host  cell  the  elementary  body  grows  in  size,  and 
becomes  an  initial  body  (fig.  119,  a), 

3.  A  reaction  on  the  part  of  the  host  cell  results,  for  nucleolar, 
plastin  substance  is  extruded  from  the  cell-nucleus  and  surrounds 
the  parasitic  initial  body.  The  latter  is  thus  enveloped  in  a  mantle 
(hence  the  name  Chlamydozoa,  from  )(Xa/jLv<;,  a  mantle),  and  the 
characteristic  cell-inclusion  (Guarnieri's  body,  Negri's  body,  etc.)  is 
produced.  The  nucleolar,  mantle  substance  probably  represents  the 
^*  cytoplasm  "  of  Cytoryctes,  described  by  Calkins. 


210  THE    ANIMAL    PARASITES    OF    MAN 

4.  The  body  next  breaks  up  into  a  number  of  smaller  bodies 
known  as  initial  corpuscles.  These,  in  their  turn,  divide  by  simple 
division  (in  the  manner  already  described)  into  numerous  elementary 
bodies  (fig.  119).     Thus,  the  life-cycle  is   completed. 

The  Chlamydozoa  are,  then,  the  minute  granules  inside  the  body  of 
the  Cytoryctes  variolce  or  the  Neuroryctes  hydrophobics,  so  that  the  whole 
body  of  the  Cytoryctes  or  Neuroryctes  corresponds  to  the  mantle  and 
parasite  of  the  Chlamydozoon.  The  Cytoryctes  group  is  said  to 
cause  destruction  of  the  host  cell.  The  Cytooikon  group  (e.g.y 
trachoma  bodies)  causes  proliferation  of  the  host  cell. 

In  September,  1913,  Noguchi  ^  described  the  cultivation  of  the 
parasite  of  rabies  in  an  artificial  medium,  similar  to  that  used  by.<him 
for  the  cultivation  of  Spirochceta  recur rentis.  The  cultures  were  stated 
to  be  infective  to  dogs,  rabbits  and  guinea-pigs.  Levaditi,  in 
December,  1913,  stated  that  he  had  succeeded  in  cultivating  spinal 
ganglia  of  rabid  monkeys  in  monkey  plasma. 

Noguchi  and  Cohen  (November,  1913)^  have  succeeded  in 
cultivating  the  so-called  trachoma  bodies,  or  at  any  rate  bodies 
very  closely  resembling  them  morphologically.  The  medium 
employed  was  Noguchi's  ascitic  fluid  and  rabbit  kidney  medium,  as 
used  for  spirochaetes.  The  coarser  cultural  forms  stained  blue  w^th 
Giemsa's  solution,  the  finer  ones  stained  red.  Attempts  to  infect 
monkeys  from  the  culture  tubes  failed. 

From  their  behaviour  on  treatment  with  such  reagents  as  saponin, 
bile  and  sodium  taurocholate,  Prowazek  considers  that  the  Chlamy- 
dozoa approach  the  Protozoa. 


PROTOZOA  INCERT^  SEDIS. 

Sergentella    hominis,    Brumpt,    1910. 

Et.  and  Ed.  Sergent  in  1908  found  vermiform  bodies  about  40  fi 
long  by  I  /[*  to  I'S  f^  broad  in  the  blood  of  an  Algerian  suffering  from 
nausea  and  cold  sweats,  without  other  symptoms.  The  bodies  were 
pointed  at  each  end,  with  a  somewhat  ill-defined  nucleus  in  the 
middle.     Their  systematic  position  is  doubtful. 


'  Journ.  Exptl.  Med..,  xviii,  p.  314. 
"^  Idem,  p.  572. 


PLATYHELMINTHES  211 


B.     PLATYHELMINTHES.  or  Flat  Worms. 

BY 

J.  W.  W.  STEPHENS,   M.D.,  B.C.,  D.P.H.' 

Definition  :  Bilaterally  symmetrical  animals  without  limbs,  the  form  of  which 
is  leaf  or  tape-like,  rarely  cylindrical,  and  whose  primary  body  cavity  (segmentation 
cavity)  is  absent,  the  cavity  being  filled  by  a  mesenchymatous  tissue  (parenchyma). 

The  mouth  is  either  situated  at  the  anterior  end  of  the  body,  or  is  shifted  more 
or  less  backwards  on  to  the  flat  ventral  surface.     The  alimentary  canal  consists  of  a 


Note. — An  Appendix  on  Protozoology  will  be 
found  on  pp.  733-752.  This  has  been  prepared  in 
order  to  incorporate  a  number  of  new  additions  to 
knowledge  made  since  the  body  of  the  book  was 
printed  off. 


To  Binder  :  face  p.   210. 


the  capillary  processes  of  which  go  on  uniting  into  larger  branches, ^nd  finally  form 
two  large  collecting  vessels,  which,  sometimes  separately  and  sometimes  united, 
open  to  the  exterior  through  one,  two,  or  numerous  pores.      'j; 

Nearly  all  the  Platyhelminthes  are  hermaphroditic,  and  in  Jiearly  all  there  are, 
in  addition  to  the  ovaries  producing  ova,  other  glands  attached  to  the  female  genital 
apparatus,  namely,  the  vitellaria  or  yolk  glands,  which  provide  a  substance  termed 
yolk,  which  serves  as  nourishment  for  the  embryo.  The  fully  formed  eggs  have 
shells  and  are  "  compound,"  z>.,  composed  of  the  egg  or  ovarian  cell,  which  is 
surrounded  by  numerous  yolk  cells  or  their  products  of  disintegration.  The  two 
sexual  openings  usually  lie  close  together,  frequently  in  the  fundus  of  a  genital 
atrium  ;  they  are  rarely  separated  from  one  another.  Shell  glands  also  usually 
occur  (p.  221). 

Reproduction  is  sexual,  often,  however,  combined  with  asexual  methods  of 
propagation  (segmentation,  budding).  The  Platyhelminthes  live  partly  free  in  fresh 
or  salt  water,  exceptionally  also  on  land.  The  greater  part,  however,  live  as 
parasites  on  or  in  animals. 


2IO  THE    ANIMAL    PARASITES    OF    MAN 

4.  The  body  next  breaks  up  into  a  number  of  smaller  bodies 
known  as  initial  corpuscles.  These,  in  their  turn,  divide  by  simple 
division  (in  the  manner  already  described)  into  numerous  elementary 
bodies  (fig.  119).     Thus,  the  life-cycle  is   completed. 

The  Chlamydozoa  are,  then,  the  minute  granules  inside  the  body  of 
the  Cytoryctes  variolce  or  the  Neuroryctes  hydropJiobice,  so  that  the  whole 
body  of  the  Cytoryctes  or  Neuroryctes  corresponds  to  the  mantle  and 
parasite  of  the  Chlamydozoon.  The  Cytoryctes  group  is  said  to 
cause  destruction  of  the  host  cell.  The  Cytooikon  group  {e.g.y 
trachoma  bodies)  causes  proliferation  of  the  host  cell. 

In  September,  1913,  Noguchi  ^  described  the  cultivation  of  the 
parasite  of  rabies  in  an  artificial  medium,  similar  to  that  used  by,<him 
for  the  cultivation  of  Spirochcvta  recur rentis.  The  cultures  were  stated 
i.*o    hf^    infprfivp    to    rlncr«;     rabbits    and    £^uinea-Di?s.       Levaditi,    in 


■^ — CTCTITT^. 

Sergentella    hominis,    Brumpt,    1910. 

Et.  and  Ed.  Sergent  in  1908  found  vermiform  bodies  about  40  ^ 
long  by  I  ft  to  1*5  yL6  broad  in  the  blood  of  an  Algerian  suffering  from 
nausea  and  cold  sweats,  without  other  symptoms.  The  bodies  were 
pointed  at  each  end,  with  a  somewhat  ill-defined  nucleus  in  the 
middle.     Their  systematic  position  is  doubtful. 


'  Joum,  Exptl.  Med.,  xviii,  p.  314. 
2  Idem,  p.  572. 


PLATYHELMINTHES  21 


B.     PLATYHELMINTHES,  or  Flat  Worms. 

BY 

J.  W.  W.  STEPHENS,   M.D.,  B.C.,  D.P.H.' 

Definition  :  Bilaterally  symmetrical  animals  without  limbs,  the  form  of  which 
is  leaf  or  tape-like,  rarely  cylindrical,  and  whose  primary  body  cavity  (segmentation 
cavity)  is  absent,  the  cavity  being  filled  by  a  mesenchymatous  tissue  (parenchyma). 

The  mouth  is  either  situated  at  the  anterior  end  of  the  body,  or  is  shifted  more 
or  less  backwards  on  to  the  flat  ventral  surface.  The  alimentary  canal  consists  of  a 
short  fore-gut,  which  is  frequently  provided  with  a  muscular  pharynx,  and  of  a  simple 
forked  or  branched  mid-gut  ;  there  is  neither  a  hind-gut  nor  an  anus  ;  in  one  class, 
the  Cestodes,  the  alimentary  canal  has  entirely  disappeared  except  for  muscular 
remnants  in  the  scolex. 

The  INTEGUMENT  OF  THE  BODY  consists  either  of  a  ciliated  epithelium  of  only 
one  layer  (Turbellaria),  or  of  a  cuticle  and  gland-like  cells  embedded  in  the 
parenchyma,  or  subcuticular  layer  (Cestodes,  Trematodes).  The  dermo-muscular 
layer  consists  of  annular,  longitudinal,  and  even  diagonal  fibres,  while  the 
parenchyma  is  traversed  by  dorso-ventral  fibres. 

The  central  nervous  system,  which  is  embedded  in  the  parenchyma  of  the 
body,  consists  of  cerebral  ganglia,  united  together  in  the  shape  of  dumb-bells,  and 
of  two  or  more  longitudinal  medullary  fascicles,  often  forming  transverse 
anastomoses.  Organs  of  sense  usually  occur  only  in  the  free-living  species,  more 
rarely  during  the  free-living  stages  of  a  few  parasitic  species  and  in  a  few  ecto>- 
parasitic  forms. 

[In  Platyhelminthes  simple  eye-spots  frequently  occur,  and  in  a  few  an  auditory 
vesicle.]  z.  . 

Blood-vessels  and  definite  respiratory  organs  are  lacking  [except  in 
Nemertinea\  ;  the  EXCRETORY  APPARATUS  (formerly  termed  water- vascular  system) 
is  typical  of  the  entire  class.  It  commences  in  the  interstices  of  the  parenchyma, 
with  peculiar  terminal  cells  (ciliated  funnels;,  which  will  be  described  later  (p.  219), 
the  capillary  processes  of  which  go  on  uniting  into  larger  branches^ ^nd  finally  form 
two  large  collecting  vessels,  which,  sometimes  separately  and.  sometimes  united, 
open  to  the  exterior  through  one,  two,  or  numerous  pores.     -^ 

Nearly  all  the  Platyhelminthes  are  hermaphroditic,  and  in  .nearly  all  there  are, 
in  addition  to  the  ovaries  producing  ova,  other  glands  attached  to  the  female  genital 
apparatus,  namely,  the  vitellaria  or  yolk  glands,  which  provide  a  substance  termed 
yolk,  which  serves  as  nourishment  for  the  embryo.  The  fully  formed  eggs  have 
shells  and  are  "  compound,"  i.e.^  composed  of  the  ^zg  or  ovarian  cell,  which  is 
surrounded  by  numerous  yolk  cells  or  their  products  of  disintegration.  The  two 
sexual  openings  usually  lie  close  together,  frequently  in  the  fundus  of  a  genital 
atrium  ;  they  are  rarely  separated  from  one  another.  Shell  glands  also  usually 
occur  (p.  221). 

Reproduction  is  sexual,  often,  however,  combined  with  asexual  methods  of 
propagation  (segmentation,  budding).  The  Platyhelminthes  live  partly  free  in  fresh 
or  salt  water,  exceptionally  also  on  land.  The  greater  part,  however,  live  as 
parasites  on  or  in  animals. 


212  the  animal  parasites  of  man 

Classification  of  the  Platyhelminthes. 

Class  /.— Turbellaria  (or  Eddy  Worms).  Flat  worms  for  the  most  part,  free 
living,  and  always  covered  with  a  ciliated  epithelium. 

Order  i. — Rhabdocoelida^  gut  unbranched. 

Order  2. — Tricladida^  gut  with  three  main  branches. 

Order  -^.—Polydadida,  a  central  gut  with  lateral  caeca.  Development  direct 
or  through  metamorphosis.  They  live  in  fresh  and  salt  water  or  on  land  ; 
very  seldom  as  parasites. 

Class  //.— Trematoda  (Sucking  Worms').  [Usually  known  as  Flukes. — F.  V.  T.] 
Flat  worms,  living  as  ecto-  or  endoparasites,  that  are  only  ciliated  in  the  larval 
condition,  and  in  their  adult  state  are  covered  with  a  cuticle,  the  matrix  cells  of 
which  lie  in  the  parenchyma.  They  have  either  one,  a  few,  or  several  suckers,- 
and  frequently  also  possess  chitinous  fixation  and  adhesive  organs.  The  intestine 
is  single,  but  generally  bifurcated,  and  not  uncommonly  there  are  transverse  anasto- 
moses between  the  forks  or  diverticula  on  them.  Excretory  organs  double,  with 
two  orifices  at  the  anterior  extremity  or  a  single  one  at  the  posterior  end. 
Development  takes  place  by  a  metamorphosis  or  alternation  of  generations  (p.  283). 
These  worms  are  almost  always  hermaphroditic,  with  two  or  more  female  and  one 
male  sexual  orifice.  They  live,  almost  without  exception,  as  parasites  on  vertebrate 
animals,  but  the  intermediate  generations  are  passed  in  molluscs. 

Class  III. — Cestoda  (Tapeworms).  Endoparasitic  flat  worms  without  an  alimen- 
tary canal.  The  larval  stages  are  rarely  ciliated,  but  are  usually  provided  with  six 
spines  ;  the  adult  worm  is  covered  with  a  cuticle,  the  matrix  cells  of  which  are  em- 
bedded in  the  parenchyma.  The  body  consists  of  a  single  segment  (Cestodaria)  or  a 
chain  of  segments,  in  which  case  it  consists  of  the  scolex  and  the  segments  contain- 
ing the  sexual  organs  (proglottides)  (Cestodes  s.  str.).  The  scolex  is  provided  with 
various  adhesive  and  fixation  organs,  and  there  are  calcareous  corpuscles  in  the 
parenchyma.  Excretory  organs  symmetrical,  opening  at  the  posterior  end.  These 
worms  are  always  hermaphroditic,  and  then  possess  one  or  two  female  and  one 
male  sexual  orifice.  During  development  a  larval  intermediate  stage  ("measle") 
occurs  and  almost  always  in  a  different  host  to  that  in  which  the  adult  sexual  worm 
lives.  The  adult  stage  is  parasitic  in  vertebrate  animals^  but  the  larval  stage  may 
occur  in  invertebrates. 

Class   II.     TREMATODA,  Rud. 

These  worms  are  usually  leaf-  or  tongue-shaped,  but  also  barrel- 
shaped  or  conical;  they  vary  from  o*i  mm.  to  almost  i  m.^  in 
length  ;  most  of  them,  however,  are  small  (5  mm.  to  15  mm.).  The  sur- 
face on  which  the  orifice  of  the  uterus  and  the  male  sexual  opening 
are  situated  is  termed  the  ventral  surface ;  the  oral  aperture,  which 
also  acts  as  anus,  is  always  at  the  anterior  end  in  the  sub-order 
Prostomata  (p.  230),  but  in  the  sub-order  Gasterostomata  it  is  ventral. 

'  This  grouping  goes  back  to  the  year  1800,  and  wai  made  by  J.  G.  H.  Zeder,  a  physician 
and  helminthologist  of  Forchheim,  who  divided  the  helminths,  which  until  1851  were  generally 
regarded  as  a  special  class  of  animals,  into  the  groups  of  round,  hook,  sucker,  tape  and 
bladder  worms,  as  which  they  are  recognized  up  to  the  present  time.  In  1809,  K.  A.  Rudolphi 
gave  them  the  names  Nematodes,  Acanthocephali,  Trematodes,  Cestodes  and  Cystici. 

2  A  sucker  or  acetabulum  (little  cup)  is  a  round,  cup-shaped  muscular  organ,  the  muscles  of 
which  are  sharply  defined  from  those  of  the  body. 

2  Netnatobothrium  filarina,  van  Bened.,  on  the  branchial  chamber  of  the  Tunny. 


TREMATODA  21 3 

Suckers  are  always  present  and  occur  in  varying  numbers  and 
positions  at  the  anterior  extremities  as  well  as  on  the  ventral  surface, 
and  occasionally  on  the  lateral  margin  and  on  the  dorsum ;  the 
beginning  of  the  intestine  (mouth)  is  always  surrounded  by  a  sucker 
in  the  Prostomata. 

In  or  near  the  suckers  there  may  be  chitinous  hooks,  claws  or 
claspers,  or  the  surface  of  the  body  is  more  or  less  covered  with  spines, 
scales  or  prickles;  in  one  genus  (Rhopalias)  there  are  projectile  tentacles 
beset  with  spines  on  the  sides  of  the  anterior  part  of  the  body. 

The  body  of  adult  Trematodes  is  covered  by  a  homogeneous  layer 
of  varying  thickness,  which  either  lies  directly  over  the  external 
layer  (basement  membrane)  of  the  parenchyma,  or  over  the  muscles 
embedded  in  the  parenchyma.  This  investing  membrane  (cuticle) 
arises  from  pear-shaped  or  spindle-shaped  cells  arranged  singly  or  in 
groups  (which  lie  between  or  internal  to  the  diagonal  muscles),  and 
is  connected  with  them  by  processes  ;  these  cells  one  may  regard  as 
epithelial  cells  which  have  sunk  down,  or  possibly  as  parenchymatous 
cells.  An  epithelium  of  one  layer  is  also  found  on  the  body  of 
young  stages,  but  it  disappears  during  growth,  and  only  occasionally 
do  its  nuclei  persist  until  adult  life.  In  its  place  we  then  find  the 
cuticle,  which,  moreover,  extends  into  all  the  body  openings  more  or 
less  deeply. 

It  is  thus  a  debatable  point  whether  the  ^'investing  layer  "  of  flukes 
is  a  cuticle — that  is,  consists  of  modified  epithelial  cells — or  whether  it  is 
a  basement  membrane,  i.e.,  compressed  and  modified  connective  tissue 
cells ;  in  this  latter  case  the  true  epidermis  and  cuticle  have  been  cast 
off.  In  the  former  case  the  epidermal  cells  are  the  pear-shaped  cells 
referred  to  above.  According  to  recent  authors  it  consists  of  two 
parts,  an  outer  true  cuticle  and  an  inner  basement  membrane.  There 
are  also  unicellular  cuticular  glands,  lying  isolated  or  in  groups,  which 
are  termed  cephalic,  abdominal,  or  dorsal  glands  according  to  the 
position  of  their  orifice. 

The  PARENCHYMA  is  a  connective  substance,  the  structure  of 
which  is  still  a  matter  of  dispute.  It  consists,  according  to  some 
authors,  of  multipolar  cells,  the  offshoots  from  which  anastomose  with 
each  other  so  that  a  network,  permeating  the  entire  body  and  encom- 
passing all  the  organs,  is  produced.  There  exists  also,  as  part  of  it,  a 
homogeneous  matrix,  in  the  form  of  lamellae  and  trabeculae  that  border 
small  cavities  communicating  with  each  other  and  filled  with  fluid. 
According  to  other  authors,  the  parenchyma  of  the  Trematodes  con- 
sisted originally  of  cells,  of  which,  however,  only  the  cell  membranes 
remain,  while  the  protoplasm  has  been  liquefied  except  for  small  residua 
around  the  nucleus.  Between  these  cells  an  intercellular  mass  has 
appeared.      By  partial  absorption  of  the  walls,  adjoining  spaces  unite. 


214 


THE   ANIMAL   PARASITES   OF   MAN 


and  the  originally  flat  cell  walls  become  transformed  into  trabeculae. 
According  to  this  vi^w  the  cavities  filled  with  fluid  are  zw/ra-cellular, 
according  to  the  former  view  /«/^y-cellular.  Pigment  cells  occur  only 
in  a  few  species. 

The  MUSCULAR  SYSTEM  of  the  Trematodes  is  composed  of  (i)  a 
dermo-muscular  tube,  (2)  the  dorso-ventral  or  parenchymal  muscles, 
(3)  the  suckers,  and  (4)  the  special  muscles  of  certain  organs. 

The  dermo-muscular  tube,  which  lies  fairly  close  to  the  cuticle, 
consists  of  annular,  diagonal,  and  longitudinal  fibres  which  surround 
the  entire  body  in  one  or  several  layers,  and  as  a  rule  are  more  strongly 
developed  on  the  ventral  surface  as  well  as  in  the  anterior  part  of  the 
body.     The  MUSCLES  OF  the  parenchyma  are  found  chiefly  in  the 

7.  I. 

A  \  Md. 

F.v.s.       C7t- 


F.v.s 


F.v.s 


Ex.v. 


Fig.  120. — Half  of  a  transverse  section  through  Fasciola  hepatica,  L.  25/1.  Cm.,  Cuticle 
with  scales;  under  the  cuticle  are  circular  muscles,  and  adjoining  them  the  longitudinal  and 
diagonal  muscles;  internal  to  the  latter  are  the  matrix  cells  of  the  cuticle;  /.,  gut;  the 
other  similarly  contoured  cavities  are  gut  diverticula  that  have  been  transversely  or  obliquely 
sectioned;  i^.z^.^.,  vitellaria  ;  ^.^.z/.,  excretory  vessels;  T.,  testes;  J/<;/. ,  median  plane;  the 
fibres  passing  from  the  ventral  to  the  dorsal  surface  are  the  muscles  of  the  parenchyma.  The 
parenchyma  itself  is  omitted. 

lateral  parts  of  the  body  and  pass  through  the  parenchyma  in  a 
dorso-ventral  direction  ;  their  diverging  brush-like  ends  are  inserted  on 
the  inner  surface  of  the  cuticle  (fig,  120). 

The  suckers  are  specially  differentiated  parts  of  the  dermo- 
muscular  tube.  Their  concave  inner  surface  is  lined  by  the  con- 
tinuation of  the  cuticle  and  their  convex  external  surface  is  covered 
by  a  more  dense  tissue  that  frequently  takes  the  form  of  a  refractive 
membrane,  thus  separating  them  from  the  parenchymal  muscles. 

The  principal  mass  of  the  suckers  consists  of  muscular  fibres  which 
run  in  three  directions  —  equatorial,  meridional  and  radial.  The 
equatorial  fibres  correspond  to  the  annular  muscles,  the  meridional 
fibres  to  the  longitudinal  muscles,  and  the  radial  fibres  to  the  muscles 
of  the  parenchyma;  the  radial   fibres  are  always  the  most  strongly 


TREMATODA 


215 


developed.  The  function  of  these  muscles  is  evident  from  their 
position  ;  the  meridional  fibres  flatten  the  suctorial  disc  and  diminish 
the  depth  of  its  cavity,  so  that  the  internal  surface  may  adhere  to 
the  object  to  be  held  ;  if  the  equatorial  fibres  now  contract,  the 
sucker  rises  by  elongating  longitudinally,  and  its  inner  surface  is 
drawn  in  by  the  contraction  of  the  radial  muscles.  Thus  the 
sucking   disc  becomes  adherent.     Usually  also  there  is  a  sphincter 


Fig.  121. — Harmostoinum  leptostomum,  Olss.,  an  immature 
specimen  from  Helix  hortensis.  Nervous  system^  according  to 
Bettendorf.  A.s.,  ventral  sucker;  Cg..,  cerebral  ganglion; 
Ex.p.,  excretory  pore  ;  G.p.,  genital  pore;  6>.^.,  oral  sucker; 
M.d.,  dorsal  medullary  nerve;  M.I.,  lateral  medullary  nerve  ; 
N.ph.,  pharyngeal  neive  ;  M.v.,  ventral  medullary  nerve. 
Magnified. 


at  the  border  of  the  suckers,  which  plays  its  part  during  the  act  of 
adhesion  by  constricting  in  a  circular  manner  that  part  of  the  mucous 
membrane  to  which  it  is  attached.  The  loosening  of  the  fixed  sucker 
is  effected  by  relaxation  chiefly  of  the  radial  fibres,  by  the  contraction 
of  the  meridional  fibres  and  certain  bundles  of  muscles  situated  at 
the  base  and  at  the  periphery  of  the  suckers.  The  connective  and 
elastic  tissues  between  the  muscles  of  the  suckers  probably  also  take 
part  in  the  process. 


2l6  THE   ANIMAL   PARASITES   OF   MAN 

Of  the  muscles  of  the  organs  which  have  developed  from  the 
parenchyma  muscles  we  may  briefly  mention  those  bundles  that  are 
attached  to  certain  parts  of  the  genital  apparatus,  to  the  suckers, 
to  the  hooks  and  claws,  and  also,  at  all  events  in  Fasciola  hepatica, 
to  the  spines.  The  sheaths  used  for  the  projection  of  the  tentacles 
of  the  RhopaliadcB  are  also  muscular. 

The  contractile  elements  consist  of  fibres  of  various  lengths 
that  are  mostly  parallel  to  one  another,  and  frequently  anastomose ; 
a  cortical  substance  finely  fibrillated  can  usually  be  distinguished 
from  an  internal  homogeneous  mass ;  large  nucleated  cells  of  uniform 
size  are  always  connected  with  them ;  these  have  been  variously 
interpreted,  but  have  been  proved  to  be  myoblasts,  one  or  more 
of  their  processes  constituting  the  muscular  fibres. 

The  MOVEMENTS  of  the  Trematodes  consist  in  alterations  of  form 
and  position  of  the  body,  as  well  as  in  creeping  movements. 

In  the  NERVOUS  system  (fig.  121)  can  be  distinguished  a  cerebral 
portion  as  well  as  strands  (medullary  strands)  running  from  it^ 
and  peripheral  nerves.  The  cerebral  portion  always  consists  of 
two  large  ganglia  situated  in  the  anterior  end  of  the  body  which 
pass  dorsally  over  the  oesophagus  and  are  connected  by  means 
of  a  broad  and  thick  commissure  composed  of  fibres  only.  From 
each  ganglion  three  nerves  run  anteriorly — the  inner  and  dorsal 
nerve  for  supplying  the  anterior  dorsal  part  of  the  body ;  the 
median  and  ventral  for  the  oral  sucker;  and  the  exterior  and  lateral 
likewise  for  the  supply  of  the  sucker. 

In  a  similar  manner  three  strands  run  backwards  from  each 
ganglion — one  dorsal,  one  lateral  and  one  ventral.  The  dorsal  and 
ventral  strands  become  united  and  curve  backwards  ;  the  symmetrical 
lateral  strands  are  connected  by  means  of  transverse  commissures, 
the  number  of  which  vary  according  to  the  species.  Such  commis- 
sures also  exist  between  the  lateral  and  the  two  other  strands  on  each 
side.  There  are  ganglion  cells  along  the  entire  course  of  the  posterior 
cords,  more  particularly  at  the  points  of  origin  of  the  commissures. 
There  also  appears  to  be  in  addition  a  fourth  anterior  and  posterior 
pair  of  nerves,  the  front  pair  for  the  oral  sucker  and  the  hind  pair 
for  the  pharynx. 

The  peripheral  nerves,  which  spring  from  the  posterior  strands 
as  well  as  from  the  commissures,  either  pass  directly  to  the  muscular 
fibres  or  to  the  sensory  cells  that  are  situated  at  the  level  of  the 
subcuticular  cells,  or  they  reach  these  after  the  formation  of  a  plexus 
situated  immediately  beneath  the  dermo-muscular  layer ;  the  pro- 
cesses directed  outwards  terminate  in  small  vesicles  in  the  cuticle. 

As  to  other  ORGANS  OF  SENSE,  simple  eyes,  two  or  four  in 
number,  are  known  in  several   ectoparasitic  species  as  well  as  in  a 


TREMATODA 


217- 


few  free-living  larval  stages  (Cercariae)  of  endoparasitic  forms.     In  the 
adult  stage,  however,  they  usually  undergo  complete  atrophy. 

The  ALIMENTARY  CANAL  commences  with  an  oral  aperture, 
generally  terminal  or  sub-terminal  (ventral)  at  the  anterior  extremity, 
which  leads  into  an  oral  cavity  usually  surrounded  by  a  sucker  ;. 
the  oesophagus,  of  various  lengths,  is  directed  backwards  and  is 
generally  surrounded  by  a  muscular  pharynx  (fig.  122).  In  some 
cases  there  exists  between  the  sucker  and  pharynx,  pharyngeal  pouches 
(praepharynx).  Sooner  or  later  the 
intestine  divides  into  two  lateral 
branches  directed  backwards,  both  of 
which  end  blindly  (caeca)  at  the  same 
level.'  In  many  ectoparasites  (Mono- 
genea  [p.  222])  a  connection  exists 
between  the  genital  glands  and  one  of 
the  intestinal  branches  (ductus  vitello- 
intestinalis  [fig.  123]). 

The  oral  cavity,  pharyngeal  pouches, 
pharynx,  and  oesophagus  are  lined  with 
a  continuation  of  the  cuticle  of  the 
body  ;  the  gut  caeca  are  lined  with  tall 
cylindrical  epithelium  (fig.  120).  The 
oesophagus  and  intestinal  branches 
often  have  also  one  layer  of  annular 
and  longitudinal  muscles;  the  pharynx 
has  essentially  the  structure  of  a  sucker 
(fig.  122). 

The    accessory  organs    of    the   ali- 
mentary canal  consist  of  groups  of  unicellular  salivary  GLANDS  that 
discharge  into  the  oesophagus  in  front  of  or  behind  the  pharynx,  or 
even  into  the  pharynx  itself. 


Fio.  122. — Median  section  through 
the  anterior  part  of  Fasciola  hepatica  : 
the  oral  sucker,  pharyngeal  pouches, 
pharynx,  oesophaeus,  cuticle  with 
spines,  and  the  body  parenchyma. 


^  The  following  conditions  represent  deviations  from  this  type:  (i)  In  Gasterostomum 
the  oral  apeiture  is  situated  in  the  middle  of  the  ventral  surface,  and  occasionally  is 
even  nearer  to  the  posterior  than  to  the  anterior  end.  There  is  no  proper  oral  sucker,, 
but  the  pharynx  is  thus  termed.  (2)  A  few  genera,  such  as  G aster ostomum^  Aspidogaster ^ 
Diplozoon,  etc.,  have  only  one  intestinal  diverticulum,  which  is  undoubtedly  to  be  taken 
as  representing  the  primitive  condition,  as  it  is  also  often  found  in  the  young  stages 
of  the  Trematoda.  (3)  The  branches  of  the  intestines  are  curved  and  united  behind 
(several  Tristomidce  and  Monostomida),  while  in  Polystomtwi  integerrimum  (in  the  bladder 
of  frogs)  there  are  several  commissures  betwee  n  the  intestinal  branches,  and  in  the  ^r-^zV/^- 
somidce  the  united  intestinal  branches  proceed  as  one  channel  towards  the  posterior  end. 
(4)  The  termination  of  the  two  intestinal  branches  is  not  always  on  a  level ;  they  are 
therefore  of  different  lengths.  (5)  When  the  oesophagus  is  very  long  the  intestinal  branches 
extend  both  forward  and  backward,  so  that  the  gut  exhibits  the  form  of  an  H.  (6)  In 
the  broad  and  fiat  species  the  gut-forks  form  diverticula  mostly  externally  but  also  internally  ; 
these  again  may  branch  (fig.  139).  (7)  In  a  few  cases  [Nematobothrium,  Didymozoon)  the 
intestine  completely  disappears  up  to  the  pharynx. 


2l8 


THE   ANIMAL   PARASITES   OF   MAN 


The  food  of  the  Trematodes  consists  of  mucus,  epithehal  cells,  the 
intestinal  contents  of  the  hosts,  and  often  also  of  blood,  and  this  not 
only  in  those  species  living  in  the  vascular  system,  but  also  in  species 


Oot, 


Ir.  c.v 


v.d.e. 


Fig.  123.  —  Polystomum  integerrimum,  a 
-monogeneiic  fluke  from  the  urinary  bladder 
of  the  frog,  z.,  intestine;  h.,  large  hooks 
of  the  sucking  disc  ;  h.k.^  smaller  hooklets  ; 
Lev.,  longitudinal  vitelline  ducts;  0.,  oral 
orifice  ;  OoL,  ooiype  ;  ov.,  ovary  ;  s.p.,  suckers 
of  the  disc  ;  tr.c.v.y  transverse  vitelline  ducts  ; 
Ut.y  uterus  with  ova;  7;.,  entrance  to  the 
vagina;  v.d.e.,  vas  deferens;  v.d.i.,  ductus 
vitello-intestinalis  ;  the  vitellaria  and  testes 
are  not  shown.     Magnified.     (After  Zeller.) 


Fig.  124. — Allocreadium  iso- 
porum,  Looss.  Excretory  appa- 
ratus. Of  the  other  organs,  the 
oral  sucker,  pharynx,  genital 
pore,  ventral  sucker,  ovary  and 
testes  are  shown ;  the  cylin- 
drical excretory  bladder  is  in 
the  posterior  end.  38/1.  (After 
Looss.) 


living  as  ectoparasites  or  in  the  intestine  or  biliary  passages  of  their 
hosts. 

The  final  products  of  assimilation  dissolved  in  the  fluids  of  the 
body   are   distributed  throughout   the   parenchyma   and   are   thence 


TREMATODA 


219 


expelled  by  a  definite  tubular  system  (excretory  apparatus,  proto- 
nephridia,  formerly  also  termed  the  water-vascular  system).  This 
system,  which  is  distributed  throughout  the  entire  body  (fig.  124),  is 
symmetrically  developed,  and,  in  the  ectoparasitic  Trematodes,  it 
opens,  right  and  left,  at  the  anterior  end  on  the  dorsal  surface  ;  in  all 
other  flukes,  however,  it  opens  singly  into  the  excretory  pore  (foramen 
caudale)  at  the  centre  of  the  posterior  border ;  in  those  cases,  however, 
where  a  sucker  is  present  at  the  posterior  end,  as  in  the  Amphistomata, 
the  excretory  pore  is  situated  on  the  dorsal  surface  close  in  front  of 
the  sucker. 

The  EXCRETORY  SYSTEM^  consists  of  several  parts  :  (i)  of  the  more 
or  less  numerous  terminal  "flame"  cells  or  funnel  cells  (figs.  124, 125); 
(2)  of  the  capillaries  ending  in  them  ;  (3)  of 
larger  vessels  receiving  the  capillaries  ;  and  (4) 
of  the  excretory  bladder.  Terminal  cells  and 
capillaries  may  be  compared  to  unicellular 
glands  with  long  excretory  ducts  ;  the  cellular 
body  (fig.  125)  is  comparatively  large,  stretched 
longitudinally,  more  rarely  transversely,  and 
provided  with  numerous  processes,  that  are 
lost  in  the  parenchyma  ;  within  is  a  conical 
cavity  (analogous  to  the  secretory  cavity  of 
unicellular  glands)  which  is  continued  directly 
into  the  structureless  capillary ;  at  its  blind 
end  is  a  bunch  of  cilia  projecting  into  the 
cavity,  and  which,  during  life,  shows  a  flicker- 
ing motion  (ciliary  flame).  The  nucleus  is 
situated  in  the  protoplasm  of  the  terminal  cell 
at  its  blind  end. 

The  entire  apparatus  thus  begins  blindly — 
ix.j  within  the  terminal  cells,  to  which  must 
be  ascribed  the  capacity  of  taking  up  from  the  fluid  that  permeates 
the  parenchyma  the  products  which  are  first  collected  into  their  own 
cavities  and  thence  excreted  by  means  of  the  capillaries  and  vessels. 

The  vessels  possess  definite  walls,  consisting  of  a  membrane  and 
a  r^ucleated  protoplasmic  layer.  They  unite  at  many  points  on  either 
side,  and  again  pass  into  other  canals  (COLLECTING  TUBES),  which 
finally,  travelling  towards  the  posterior  end,  discharge  into  the 
excretory  bladder  (fig.  124). 

The  form  and  size  of  the  bladder  vary  much  according  to  the 
different  species,  but  it  always  possesses  its  own  flattened  epithelium, 
surrounded  by  circular  and  longitudinal  muscles,  the  circular  muscles 


Fig.  125.— Terminal  flame 
cell  of  the  excretory  system. 
n.,  nucleus  of  cell ;  c. ,  bundle 
ofcilia  forming  the  "flame"; 
p.,  processes  of  cell  extending 
into  parenchyma;  J.,  excre- 
tory capillary.     (Stephens.) 


The  following  description  relates  in  the  main  to  the  Distomata. 


220 


THE   ANIMAL   PARASITES   OF   MAN 


forming  a  sphincter  around  the  opening.  Frequently  also  the 
structure  of  the  bladder  extends  to  the  tubules  discharging  into 
it,  which  therefore  are  not  to  be  regarded  as  separate  *'  vessels," 
but  rather  as  tubular  diverticula  of  the  bladder,  directed  anteriorly. 
In  some  few  species  the  diverticula  also  branch  and  the  branches 
anastomose,  so  that  a  network  of  tubules  ensues  which  receives  the 
vessels  or  capillaries.  In  such  cases  there  are  also  ciliary  tracts  in 
the  tubules. 

The  contents  of  the  entire  appar- 
atus usually  consist  of  a  clear  or  some- 
times reddish  fluid ;  in  some  species 
there  are  larger  or  smaller  granules,  and 
occasionally  also  concretions  occur. 


vag. 


-ut. 


J^ec.  ut. 


^ec.  sent. 


Vit.    R. 


Fig.  126. — Diagram  of  female  genitalia.     Ov.^ 


Fig.   127.— Diagram  of  male  and  part 
of  female   genitalia.      tU.,   uterus;    vag.^ 


ovary;     ovd.^     oviduct;    Z.r.,    Laurer's    canal;      vagina;    5  ,  opening  of  vagina  ;  ^.j.,  geni- 


Rec.  sem.y  receptaculum  seminis ;    Vit.  R.,  vitel- 
larian  reservoir  ;  t.v.d.,  transverse  vitelline  duct 


tal  sinus  ;   g.p.,  genital  pore;    (^  ,  opening 
of  ejaculatory  duct  or  vas   deferens ;   c.s., 


Oo.,   ootype  ;    Sh.  gl.,   shell   gland;    Rec.    ut,,      cirrus  sac;  f.,  cirrus ;/./.,  pars  prostatica ; 


receptaculum  uterinum  ;  ut.,  uterus.    (The  various 
parts  are  not  to  the  same  scale.)     (Stephens.) 


s.v.,  seminal  vesicle;  e.J.,  ejaculatory  duct 
or  vas  deferens  ;  v.e.,  vas  efferens ;  t.,  testis. 
(Stephens.) 

Sexual  Organs. — Nearly  all  the  Trematodes  are  hermaphrodites, 
and  only  a  few  (Schistosotnidce,  Koellikeria)  are  sexually  differen- 
tiated. The  sexual  organs  usually  lie  in  the  "  central  field  "  limited 
by  the  gut  caeca  ;  the  vitellaria,  on  the  other  hand,  are,  as  a  rule, 
external  to  the  gut  caeca  in  the  *'  lateral  fields." 

The  male  apparatus^  is  composed  of  two  variously  formed 
testes  (fig.  127)  (globular,  oval,  indented,  lobed,  or  ramified),  which 
may  lie  side  by  side  or  one  behind  the  other ;  from  each  testicle 
a  tube  (vas  efferens)  originates  ;  sooner  or  later,  both  tubes  as  a  rule 
unite  to  form  the  ejaculatory  duct  or  vas  deferens,  which  is  frequently 


The  following  description  relates  mainly  to  the  Distoniata. 


TREMATODA  221 

enclosed  in  a  muscular  CIRRUS  SAC,  or  more  rarely  passes  directly  into 
the  genital  pore.  The  cirrus,  which  is  the  thick  muscular  terminal 
portion  of  the  vas  deferens,  can  be  everted  and  protruded  from  the 
cirrus  sac  and  serves  as  an  organ  of  copulation.  The  walls  of  the 
muscular  portion  of  the  tube  (the  cirrus)  are  attached  to  the  walls  of 
the  cirrus  sac,  and  hence  when  the  sac  contracts  the  cirrus  cannot 
be  protruded  except  by  evagination  of  its  lumen.  Opening  into  the 
middle  portion  of  the  vas  deferens,  and  as  a  rule  enclosed  in  the  cirrus 
sac,  is  found  a  mass  of  unicellular  glands  (prostate),  the  vesicula 
seminalis  (which  is  likewise  within,  or  may  also  be  outside  the  sac) 
being  the  dilated  first  portion  of  the  vas. 

The  female  genitalia  (fig.  1 26)  consist  of  an  ovary,  usually  situated 
in  front  of  the  testes,  the  form  of  which  varies  according  to  the 
species,  the  usually  double  vitellaria,  the  ducts  and  a  number  of 
auxiliary  organs ;  the  short  oviduct  directed  towards  the  centre 
arises  from  the  ovary,  and  is  connected  in  the  median  line  with 
the  excretory  duct  of  the  vitelline  glands.  These  grape-like  glands 
possess  longitudinal  excretory  ducts,  which  assume  a  transverse 
direction  behind  the  ovary,  unite  together  at  the  median  line  and 
form  a  single  duct,  often  dilated  into  a  vitelline  receptacle,  that  unites 
with  the  oviduct.  Near  this  point,  moreover,  there  frequently  opens 
a  canal  (Laurer's  canal)  which  begins  on  the  dorsal  surface,  and 
on  the  inner  end  of  which  a  vesicle  filled  with  sperm  (receptaculum 
seminis)  usually  occurs  (fig.  126).  Moreover,  there  are  also  numerous 
radial  unicellular  glands  (shell  glands)  at  or  beyond  the  point  of 
junction  of  the  oviduct,  vitelline  ducts  and  Laurer's  canal.  In  this 
portion  of  the  duct  (ootype),  which  is  usually  dilated,  the  ovarian 
cells  are  fertilized,  surrounded  with  yolk  cells  and  shell  material, 
and  as  ova  with  shells  they  pass  into  the  uterus  (a  direct  continua- 
tion of  the  oviduct),  which,  with  its  many  convolutions,  occupies  a 
larger  or  smaller  portion  of  the  central  field,  and  runs  either  direct 
to  the  genital  pore  or,  forming  convolutions,  first  runs  posteriorly 
and  then  bends  forward  (descending  and  ascending  limbs).  In  both 
cases  the  terminal  part  lies  beside  the  cirrus  pouch  and  discharges 
beside  the  male  orifice  either  on  the  surface  of  the  body  or  into  a 
genital  atrium.  The  terminal  portion  of  the  uterus,  which  is  often 
of  a  particular  structure,  serves  as  a  vagina  (METRATERM). 

The  cirrus  sac  may  include  (i)  the  genital  atrium  {i.e.,  the 
common  sinus,  into  which  the  vas  deferens  and  vagina  may  open), 
or  (2)  a  variable  extent  of  the  vas  from  cirrus  to  seminal  vesicle.  Thus 
the  latter  may  be  outside  the  sac.  In  the  absence  of  a  sac,  the  genital 
sinus  may  be  surrounded  by  a  pseudo-sucker,  as  in  Heterophyes  (in 
some  cases  the  ventral  sucker  itself,  from  its  close  proximity  to  the 
genital  pore,  serves  as  an  accessory  copulatory  organ).     In  other  cases 


222  THE   ANIMAL    PARASITES   OF    MAN 

copulatory  organs  are  formed  by  hooks  projecting  into  the  lumen 
of  the  terminal  portion  of  the  vas. 

The  GENITAL  PORE,  which  is  the  opening  from  the  genital  sinus  on 
to  the  surface,  is  generally  situated  at  or  near  to  the  median  line  on 
the  ventral  surface  and  in  the  anterior  region  of  the  body  ;  in  most 
of  the  Distomata  it  is  in  front  of  the  ventral  sucker,  in  other  cases, 
e.g.y  in  the  Cryptocotylince,  it  is  behind.^ 

The  spermatozoa  do  not  differ  essentially  in  their  structure  from 
those  of  other  animals  ;  the  ovarian  or  Qgg  cells  are  cells  without 
integument  and  contain  a  large  nucleus  and  a  little  protoplasm;  the 
vitellaria  also  produce  nucleated  cells,  in  the  plasm  of  which  there 
are  numerous  yellow  yolk  granules  ;  the  yolk  cells  detach  themselves, 
like  the  ovarian  cells,  from  the  ovarium,  and  pass  into  the  oviduct 
to  surround  each  ovarian  cell  in  the  ootype.  They  disintegrate  sooner 
or  later  in  the  completely  formed  Qgg  and  are  utilized  as  food  by  the 
developing   embryo. 

Development  of  the  Trematodes. 

(i)  Copulation. — Observation  has  demonstrated  that  the  one  or 
two  vaginae  occurring  in  the  ectoparasitic  Trematodes  are  utilized 
as  female  organs  of  copulation,  and  that  the  copulation  is  cross  ; 
it  is  also  known  that  Laurer's  canal,  which  was  formerly  generally 
regarded  as  the  vagina,  has  only  quite  exceptionally,  if  at  all,  served 
the  digenetic  Trematodes  as  such — it  appears  to  be  homologous  with 
the  canalis  vitello-intestinalis  of  the  Monogenea? — but  the  terminal 
portion  of  the  uterus,  termed  the  metraterm,  is  used  for  copulation. 
Cross-copulation  occurs  as  well  as  auto-copulation  and  auto-fecunda- 
tion. The  spermatozoa  subsequently  pass  through  the  entire  uterus, 
which  is  still  quite  short  at  the  time  the  male  organs  are  matured ;  the 
maturation  of  which,  as  usually  is  the  case  in  hermaphrodites,  precedes 

^  The  typical  position  of  the  genitalia  is  subject  to  many  deviations,  which  are  of 
importance  in  the  differentiation  of  the  genera  and  families.  The  following  are  some  few 
of  these  deviations  :  (i)  The  genital  pore  remains  on  the  ventral  surface,  but  is  situated 
beside  or  behind  the  ventral  sucker,  or  it  becomes  marginal,  and  is  then  found  in  front  of  or 
beside  the  oral  sucker,  or  at  a  lateral  edge,  or,  finally,  in  the  centre  of  the  posterior  border  ; 
the  ducts  also  correspondingly  alter  their  direction.  (2)  The  ovary  usually  lies  in  front  of 
the  testes,  not  rarely,  however,  behind  them  or  between  them.  (3)  The  three  genital  glands 
mostly  lie  together  close  in  front  of,  or  behind,  the  centre  of  the  body ;  they  may  be  moved 
far  back,  and  may  incidentally  become  separated  one  from  the  other.  {4)  The  vitellarium 
may  be  single,  in  which  case  it  then  may  lie  in  the  central  field.  (5)  A  few  forms  possess 
but  one,  others  several  or  numerous  testes.  Amongst  the  ectoparasitic  trematodes  there  are 
also  species  with  but  one  testis  ;  but  they  mostly  have  several.  As  a  rule,  their  uterus  is  short, 
but  the  ootype  well  developed.  Special  canals  (vagina),  single  or  double,  are  used  for  copula- 
tion, not  the  uterus.  The  vitelline  ducts  also  communicate  with  the  intestine  through  the  canalis 
vitello-intestinalis  (fig.  123). 

2  Monogenea  :  Trematoda  in  which  the  anterior  sucker,  if  present,  is  double.  Development 
without  an  intermediate  host. 


TREMATODES :  DEVELOPMENT 


223 


that  of  the  female  organs.  It  is  only  later  with  the  onset  of  egg  forma- 
tion that  the  uterus  is  fully  developed.  Copulation,  however,  takes  place 
also  in  the  case  of  fully  grown  forms  with  completely  developed  uteri. 

(2)  Formation  of  the  Ova. — The  ovarian  cells  arising  from  the  ovary 
first  become  mature  after  their  entry  into  the  ootype  by  the  formation 
of  three  polar  bodies,  fertilization  then  taking  place.  At  the  same 
time  as  the  ovarian  cell  a  number  of  yolk  cells  from  the  vitellarium 
and  secretion,  drop  by  drop,  from  the  shell  gland  reach  the  ootype.^ 
The  shell  is  then  formed  during  the  generally  active  contractions  of 
the  ootype  w^alls  and  then  passes  on  into  the  uterus.  In  the  uterus  of 
the  endoparasitic  trematodes  the  eggs  accumulate  more  and  more, 
often  in  large  quantities,  while  in  ectoparasitic  species  generally  only 


Fig.  128. — Ovum  of  Fasciola 
hepatica,  L.,  cut  longitudinally. 
The  lid  has  been  lifted  in  the 
process.  Within  the  egg  are 
numerous  yolk  cells,  and  at  the 
lid  end  there  is  the  still  unseg- 
mented  ovum  (dark).     240/1. 


Fig.  129.  —  Miracidium  of 
Fasciola  hepatica  that  has  just 
hatched  from  the  egg,  with  a  dis- 
tinct cuticular  ciliatedepithelium. 
Magnified.     (From  Leuckart.) 


one  or  some  few  eggs  can  be  found.  The  completed  ova  are  of 
various  forms  and  sizes.  They  are  mostly  oval,  at  all  events  in  the 
digenetic  trematodes,  and  the  yellowish  or  brown  shell  is  provided 
with  an  opening  at  one  pole  which  is  closed  by  a  watch-glass-shaped 
lid  (operculum).  Appendages  (filaments)  on  the  shell — at  one  or  both 
poles — are  uncommon,  but  are  the  rule  in  the  ova  of  the  Monogenea 
(ectoparasitic  species). 

(3)  Deposition    of    the    Ova. — Soon    after     their     formation,     the 
Monogenea    (ectoparasitic   trematodes)    deposit    round    the    place    of 


'  [Recent  work  {e.^.,  Goldschmidt,  Zool.  Anzeiger^  xxxiv,  p.  482)  has  shown  that  the 
older  views  regarding  the  formation  of  the  egg  must  be  modified.  In  certain  species,  at  any 
rate,  the  shell  material  is  formed  by  the  yellow  droplets  of  the  yolk  glands  and  not  by  the  so- 
called  shell  gland  (Mehli's  gland)  secretion,  which  is  clear  and  watery.  The  function  of  this 
secretion  accordingly  still  requires  explanation  ;  according  to  Looss  it  serves  as  a  covering 
secretion  for  the  egg-shell  proper.  It  appears  also  that  other  granules,  the  yolk  granules  as 
distinct  from  the  shell  drop  granules,  are  not  always  used  up  during  the  development  of  the 
embryo  and  hence  do  not  function  as  yolk,  so  these  also  when  they  exist,  and  frequently 
they  are  wanting,  must  serve  some  other  purpose,  possibly  that  of  imbibing  water  for  the  use, 
of  the  embryo.— J.  W.  W.  S.] 


224  THE   ANIMAL   PARASITES   OF   MAN 

their  attachment  on  the  skin  or  the  gills  or  other  organs  of  their 
hosts,  eggs  which  attach  themselves  by  means  of  their  filaments. 
The  embryonic  development  thus  takes  place  outside  the  parent. 
This  also  holds  good  for  the  eggs  of  many  endoparasitic  species, 
although  as  a  rule  in  these  the  eggs  are  always  retained  for  a 
longer  time  in  the  uterus.  Moreover,  they  usually  here  undergo  a 
part  or  a  whole  of  their  development,  and  are  eventually  deposited 
in  those  organs  in  which  the  adult  forms  are  parasitic,  but  this  is 
not  always  the  case,  as  the  egg,  e.g.,  of  F.  hepatica  appears  in  bile  (and 
faeces)  quite  unchanged.  By  the  natural  passages  they  eventually  get 
out  of  the  body,  and  in  cases  where  such  do  not  exist,  as  in  the  case 
of  the  blood-vessels,  the  eggs  pass  out  by  means  of  the  kidneys. 

(4)  The  embryonic  developinent,  after  irregular  segmentation  of  the 
ovum  into  a  number  of  blastomeres,  leads  to  the  formation  of  a  solid 
blastosphere  or  morula,  which  is  surrounded  by  a  cellular  investing 
membrane  (yolk  envelope),  while  the  principal  mass  of  the  cells  forms 
the  embryo,  which  uses  for  its  nourishment  the  yolk  cells,  which  have 
in  the  meantime  disintegrated  {cf.  footnote,  p.  223).  Usually,  after 
the  ova  have  reached  water  the  embryos  hatch  out,  leaving  the  yolk 
envelope  in  the  egg-shell  ;  in  other  cases,  however,  the  embryos  only 
hatch  out  after  having  been  subjected  to  the  influence  of  the  intestinal 
juices,  that  is  to  say,  in  the  intestine  of  an  intermediate  host  which  has 
ingested  with  its  food  the  ova  that  have  escaped  from  the  primary  host. 

(5)  The  post-embryonic  development  of  the  Trematodes  is  accom- 
plished in  various  ways ;  the  process  is  the  most  simple  in  the  ecto- 
parasitic  species  (Monogenea),  the  young  of  which  should  certainly 
be  regarded  as  larvae,  because  they  possess  characteristics  (cilia, 
simple  gut,  etc.)  that  are  lacking  in  the  adult  worms,  but  which, 
nevertheless,  pass  into  the  adult  state  direct  after  a  relatively  simple 
metamorphosis.  In  the  Holostomata,^  a  group  found  chiefly  in  the 
intestine  of  aquatic  birds,  and  which  rarely  occur  in  other  vertebrates, 
the  ova  develop  in  water.  The  young  are  ciliated  all  over,  and,  after 
having  entered  an  intermediate  host  (leeches,  molluscs,  arthropods, 
amphibians,  fishes)  living  in  the  water,  they  undergo  a  metamor- 
phosis into  a  second  larval  stage  ;  they  then  encyst  and  await  trans- 
mission into  the  final  host,  where  they  become  adult  Metastatic 
trematodes,  i.e.,  trematodes  without  asexually  produced  generations 
(p.  229). 

In  the  remaining  so-called  digenetic  trematodes  (p.  230)  one  or  two 
asexual  generations  interpose  between  the  miracidium  and  terminal 
stage,  so  that  quite  a  number  of  adult  worms  may  originate  from 
one  egg.     Usually  the  young,  which  are  termed  miracidia^  (fig.  129), 

•  Holostomata :  Prostomata  with  (in  addition  to  the  oral  and  ventral  suckers)  a  third 
fixation  apparatus,  generally  on  a  separate  part  of  the  body. 
^   [Also  known  as  ciliated  embryos. — F.  V.  T.] 


TREMATODES :  DEVELOPMENT  225 

hatch  in  water,  where  they  move  with  the  aid  of  their  ciha.  Sooner 
or  later  they  penetrate  into  an  intermediate  host,  which  is  always 
a  snail  or  a  mussel,  and  while  certain  of  their  organs  disappear, 
they  grow  into  a  gutless  germinal  tube  (sporocyst,  fig.  131).  These 
are  simple  elongated  sacs  with  a  central  body  cavity.  They  may  or 
may  not  have  excretory  tubules.  In  these,  according  to  the  species, 
the  larval  stages  (CERCARi^)  that  will  ultimately  become  adult  worms 
are  produced,  or  another  intermediate  generation  is  first  formed,  viz.^ 
that  of  the  REDI^^  (figs.  132,  133),  which  are  always  provided  with  an 
intestine,  and  these  then  give  rise  to  cercariae  (figs.  130,  134).  The 
cercariae,  as  a  rule,  leave  their  host  and  move  about  in  the  water 
with  the  assistance  of  their  rudder-like  tails.  After  a  little  time,  how- 
ever, they  usually  again  invade  an  aquatic  animal  (worms,  molluscs, 
arthropods,  fishes,  amphibians),  then  they  lose  their  tails  and  become 


Fig.  130. — A  group  of  cercarioe  of  Echinostoma  sp.  (from 
fresh  water).     25/1. 

encysted  (fig.  135)  ;  here  they  wait  until  they  attain,  together  with 
their  host,  the  suitable  terminal  host,  and  in  this  new  situation  they 
establish  themselves  and  reach  maturity.  Or,  again,  the  cercariae  may 
themselves  encyst  in  water  or  on  foreign  bodies  (plants)  and  wait  until 
they  are  taken  up  directly  by  the  terminal  host,  e.g.y  sheep. 

Accordingly  the  following  conditions  are  necessary  for  the  com- 
pletion of  the  entire  development :  (i)  The  terminal  host  in  which 
the  adult  stage  lives ;  (2)  an  intermediate  host  into  which  the 
miracidia  penetrate  and  in  which  they  become  sporocysts ;  (3)  a 
second  intermediate  host  in  which  the  cercariae  become  encysted. 
In  certain  species,  as  in  Fasciola  hepatica,  this  second  host  is  omitted, 
as  the  cercariae  spontaneously  encyst  on  plants,  or  again  (in  other 
species)  encystment  may  occur  within  the  first  intermediate  host,  when, 

*  [In  Fasciola  hepatica  in  the  summer  months  the  rediae  give  rise  to  daughter  redise,  which 
then  give  rise  to  cercariae. — J.  W.  W.  S.] 

15 


220 


THE   ANIMAL   PARASITES   OF   MAN 


ill  fact,  the  cercarice  (which  in  this  case  do  not  acquire  an  oar-like  tail) 
do  not  swarm  out  of,  but  encyst  themselves  w^ithin  their  sporocysts. 
The  development,  moreover,  may  be  further  complicated  by  rediae 
appearing  in  addition  to  the  sporocysts,  though  this  occurs  in  the 
first  intermediate  host  and  not  in  a  second  one. 

Animals  that  harbour  adult  digenetic  Trematodes  thus  become 
infected  by  ingesting  encysted  cercariae,  which  either  occur  (i)  in 
certain  animals  (second  intermediate  hosts)  on  which  they  feed,  or 
(2)  in  water,  or  (3)  on  plants,  or  finally  (4)  in  the  first  intermediate 
host ;  whereas  animals  harbouring  encysted  cercariae  have  been  directly 
infected  by  the  corresponding  tailed  stage,  and  animals  harbouring 
germinal  tubes  (sporocysts  or  rediae)  have  been  infected  by  the 
miracidia. 

Thus  certain  species  of  ducks  and  geese  become  infected  with 
Echinostoma  echinatiim  by  devouring  certain  water-snails  {LimncBus, 


Fig.  131. — Development  of  Fasciola  hepatica,  L.  a,  the  miiacidium  in  optical  section 
showing  cephalic  lobe,  X-shaped  eye-spot  resting  on  the  cerebral  ganglion,  two  germ  balls ; 
below  each  of  these  a  flame  cell,  and  still  lower  germ  cells  lying  in  a  caviiy  (primitive  body 
cavity).  ^,  young  sporocyst  with  two  eye-spots,  and  germ  balls  ;  the  cells  lining  the  cavity 
are  not  shown.     ^,    older  sporocyst  with  a  young  redia.     Magnified.     (After  Leuckart.) 

Paludina)  in  which  the  encysted  cercariae  occur.  Oxen  become  infected 
with  Paraniphistomum  cervi  (=  Aniphistomiun  coniciim)  by  swallowing 
with  water,  cysts  of  this  species  which  occur  at  the  bottom  of  puddles 
and  pits.  Sheep  are  infected  with  Fasciola  hepatica  by  eating  grass 
to  which  the  encysted  cercariae  of  the  liver-fluke  are  attached;  our 
song-birds  infect  themselves  or  their  young  wnth  Urogoninnts  macro- 
stomus  by  tearing  off  pieces  containing  the  corresponding  sporocysts 
which  are  full  of  encysted  cercariae  from  snails  {Succinea  amphibia), 
which  act  as  the  first  intermediate  hosts,  and  eating,  or  offering  their 
young  these  pieces. 

(i)  The   MIRACIDIA   of    the   digenetic   Trematodes   are    compara- 
tively highly  organized,  and  the  mode  of  their  formation  from  the 


TREMATODES :  DEVELOPMENT 


227 


segmentation  cells  of  the  ovum  is  only  imperfectly  known.  They  have 
a  cuticular  epithelium  (fig.  129)  entirely  or  partly  covered  with  cilia, 
beneath  this  a  dermo-muscular  tube  composed  of  circular  and  longi- 
tudinal muscles ;  also,  a  simple  gut  sac  with  an  oesophagus,  occa- 
sionally also  with  pharynx,  salivary  glands  and  boring  spine,  also 
a  cerebral  ganglion  on  which,  in  some  species,  there  are  eyes 
(fig.  131,  <^).  As  to  the  excretory  organs,  they  are  represented  by  two 
symmetrically  placed  terminal  flame  cells,  with  excretory  vessels 
opening  separately ;  there  is  a 
more  or  less  ample  (primary)  body 
cavity  between  the  parietes  of  the 
body  and  the  gut  ;  from  the  cel- 
lular parietal  lining  of  this  cavity 
single  cells  (germ  cells)  become 
free  (fig.  131,  a,  6),  and  become 
rediae  or  cercariae. 

[The  germ  cells  of  the  mira- 
cidium  and  the  germ  balls  of  the 
sporocyst  arise,  according  to  some 
observers,  by  further  division  of 
undifferentiated  blastomeres  ;  ac- 
cording to  others  from  the  cells  of 
the  lining  wall  of  its  body  cavity. 
It  is  from  these  free  germ  balls 
that  the  redia  stage  is  developed. 

[In  the  germ  ball  or  morula 
appears  an  invagination,  giving 
rise  to  the  cup-shaped  gastrula 
stage.  This  elongates  and  forms 
the  REDIA  (fig.  131,  c). 

[In  the  interior  of  the  redia 
cells  are  budded  off  and  develop 
into  gastrulae,  as  in  the  case  of  the 
sporocyst.  These  become  a  fresh 
generation  of  rediae  or  give  rise 
to  the  third  stage  (cercaria).] 

(2)  The  SPOROCYSTS,  on  the  contrary,  which  are  produced  direct 
from  the  miracidia,  are  very  simple,  as  all  the  organs  of  the  latter 
disappear,  even  to  the  muscles  and  excretory  organs,  during  or  after 
penetration  into  the  intermediate  host,  whereas  the  budded  and  still 
budding  cells  of  the  wall  of  the  (primary)  body  cavity  continue  to 
develop  rapidly  and  form  germ  balls.  The  sporocysts  when  fully 
developed  have  the  appearance  of  tubes  or  fusiform  bodies  with 
rounded  edge;  they  are  frequently  of  a  yellow  colour.     Their  length 


Fig.  132. — Young 
redia  of  Fasciola  he- 
patica^  with  pharynx 
and  intestine,  with  a 
circular  ridge  an- 
teriorly and  a  pair 
of  processes  poste- 
riorly and  masses  of 
cells  (germ  balls)  in 
the  interior.  Mag- 
nified.  (From 
Leuckart.) 


Fig.  133.— Older 
redia  of  Distoma 
echinatum,  with  ru- 
dimentary intestine 
i. ;  cercariae,  c. ;  germ 
balls,  b. ;  and  birth 
pore,  g.     Magnified. 


228 


THE  ANIMAL   PARASITES   OF   MAX 


rarely  exceeds  a  few  millimetres ;  in  some  species  their  size  increases 
exceedingly  through  proliferation,  and  they  then  occupy  a  large  portion 
of  the  body  of  the  intermediate  host. 

(3)  The  REDiJE  (figs.  132,  133),  on  the  other 
hand,  are  more  cylindrical  and  always  have  a 
simple  intestine  of  varying  length,  provided 
with  a  pharynx  ;  they  likewise  possess,  situated 
near  the  circular  ridge,  a  ''  birth  pore  "  which 
serves  for  the  exit  of  the  cercariae  originating 
within  them. 

(4)  The  CERCARi^^  are  veiy  different;  typi- 
cally they  consist  of  the  anterior  body  and  the 
oar-like  tail  at  the  posterior  end  (fig.  134).  The 
former,  even  to  the  genitalia,  has  the  organiza- 
tion of  the  adult  digenetic  Trematodes,  and 
thus  allows  the  easy  recognition  of  at  least  the 
characters  of  that  large  group  to  which  the 
species  in  question  belongs.  On  the  other 
hand,  however,  there  are  also  organs  that  are 
lacking  in  the  adult  form,  such  as,  in  many,  the 
boring  spine  in  the  oral  sucker,  or  the  eyes 
situated  on  the  cerebral  ganglion  ;  moreover, 
also,  cutaneous  glands  (fig.  134),  the  secretion  of 
which  forms  the  cyst  membrane.  The  oar-like 
tail  may  be  long  or  short  (stumpy-tailed  cer- 
caria)  or  entirely  absent ;  its  free  end  may  be 
partly  split  (furcate  cercaria),  or  split  to  its  base 
(bttcephalns)  ;  in  various  forms  also  the  an- 
terior end  of  the  tail  is  hollow,  and  has  enclosed 
within  it  the  anterior  body,  which  is  otherwise 
free.  The  size  also  of  the  cercaria  belonging 
to  the  different  species  is  very  diverse ;  in 
addition  to  forms  swimming  in  the  water  that 
have  the  appearance  of  minute  milky-white 
bodies,  there  are  forms  which  measure  as  much 
as  6  mm.  in  length. 

Fig.  135.— Encysted  cer-  °  .  1    ,      , 

caria  of  FasaWa  hepatica.  The  cncysted  ccrcariae  (fig.  135)  are  globular 

Magnified.     (After  Leuck-     qj-  oval,  and  are  surrounded  by  a  homogeneous 

art.)  ^  ^ 


Fig.  134. — Cercaria  of 
Fasciola  hepatica  ;  the  cuta- 
neous glands  at  the  side  of 
the  anterior  body.  Mag- 
nified.    (After  Leuckart.)ij^ 


^  The  cercaria  is  the  characteristic  larval  stage  of  the  Trematodes,  and  corresponds  to  a 
cysticercus  or  cysticercoid,  though  there  is  the  important  difference  that  the  cercaria  has  an 
enteric  cavity.  According  to  some  observers  the  enteron  is  represented  by  the  frontal  sucker 
of  some  Cestodes,  and  by  the  rostellum  of  the  majority  of  others. 

The  sporocyst  and  redia  are  regarded  as  intercalated  stages,  viz.,  as  cercarise  exhibiting 
pedogenesis,  i.e.,  development  of  young  by  a  parthenogenetic  process  fr(  m  individuals  [i.e.y 
cercaria)  not  yet  adult. 


TREMATODES  :     BIOLOGY  229 

membrane,  which  may  be  striated  or  contain  granules.  The  tail  is 
always  cast  off  when  encystment  occurs,  and  organs  peculiar  to  the 
cercaria  stage  (boring  papilla,  eyes)  almost  entirely  disappear.  On 
the  other  hand,  the  genitalia  appear  or  become  more  or  less  highly 
developed,  in  extreme  cases  to  such  an  extent  that  they  become 
functional,  and  after  autocopulation  the  creatures  produce  ova  within 
the  cysts. 

The  cycle  of  development  of  the  digenetic  Trematodes  has  hitherto 
been  generally  explained  as  a  typical  alternation  of  generations, 
one  sexual  generation  regularly  alternating  with  one  or  two  asexually 
reproducing  generations.  Recent  authors,  how^ever,  regard  the  cells 
in  the  sporocysts  from  which  rediae  or  eventually  cercariae  arise  as 
parthenogenetically  developing  ova,  and  the  sporocysts  as  well  as  the 
rediae  as  generations  propagating  parthenogenetically.  In  this  case, 
however,  it  is  an  alternation  of  a  sexual  not  with  an  asexual  but  with 
firstly  a  parthenogenetic  generation  (the  sporocyst),  the  central  cells 
of  which  are  regarded  as  ova  which  develop  parthenogenetically  into 
the  redia,  and  this  the  second  parthenogenetic  generation  finally 
produces  larvae  (cercariae)  capable  of  developing  into  the  sexually 
mature  form. 

Other  authors,  again,  regard  the  development  of  the  Digenea  as 
only  a  complicated  metamorphosis  (p.  283),  which  is  distributed  over 
several  generations  before  it  is  concluded. 

Biology. 

Endoparasitic  Trematodes,  as  fully  developed  organisms,  occur  in 
vertebrate  animals  only,  with  very  few  exceptions;  they  inhabit  almost 
all  the  organs  (with  the  exception  of  the  nervous  and  osseous  systems 
and  the  male  genitalia),  but  by  preference  the  intestine  in  all  its 
extent  from  the  oral  cavity  to  the  anus;  and,  further,  certain  species 
or  groups  inhabit  only  quite  restricted  parts  of  the  intestine.  Besides 
in  the  intestine  other  species  live  in  the  liver,  or  in  the  bile-ducts, 
or  in  the  gall-bladder;  other  accessory  organs  of  the  intestine,  such 
as  the  pancreas,  bursa  Fabricii  (of  birds),  are  only  infected  by  a  few 
species.  Many  inhabit  the  lungs,  or  the  air  sacs  in  fowls,  a  few 
the  trachea.  Trematodes  have  also  been  known  to  occur  in  the 
urinary  bladder,  the  urethra  and  the  kidneys  of  all  classes  of 
vertebrates;  they  are  also  present  in  the  vascular  system  of  a  few 
tortoises,  birds  and  mammals ;  in  birds  they  even  penetrate  from 
the  cloaca  into  the  oviducts,  and  are  occasionally  found  enclosed  in 
the  laid  eggs ;  one  species  is  known  to  occur  in  the  cavum 
tympani  and  in  the  Eustachian  tube  of  a  mammal  (Dugong), 
another  in  the  frontal  sinus  of  the  polecat ;  several  species  infest  the 


230  THE   ANIMAL   PARASITES   OF   MAN 

conjunctival  sac  under  the  membrana  nictitans  of  birds,  one  species 
even  lives  in  cysts  in  the  skin  of  song-birds.  In  an  analogous  manner 
the  ectoparasitic  Trematodes  are  not  entirely  confined  to  the  surface  of 
the  body  or  the  trachea  of  the  lower  vertebrate  animals ;  a  few  species 
appear  exclusively  in  the  urinafy  bladder,  in  the  oesophagus,  and  in  the 
case  of  sharks  in  an  accessory  gland  of  the  rectum. 

Trematodes  live  free  and  active  within  the  organs  attacked, 
though  they  may  attach  themselves  by  suction  for  a  longer  or  shorter 
period ;  in  other  cases,  however,  they  bore  more  or  less  deeply 
into  the  intestinal  wall  with  their  anterior  end,  or  lie  in  cysts 
of  the  intestinal  wall  which  only  communicate  with  the  lumen 
through  a  small  opening ;  in  those  species  living  in  the  lungs  of 
mammals  the  host  likewise  produces  a  cyst,  which  usually  encloses 
two  specimens;  such  association  of  a  pair  is  also  observed  in 
other  situations,  and,  though  this  is  the  rule  in  species  sexually 
distinct,  it  is  not  entirely  confined  to  these. 

As  regards  the  AGE  attained  by  endoparasitic  Trematodes,  there 
are  but  few  reliable  records,  and  these  differ  considerably ;  the 
overwhelming  majority  of  species  certainly  live  about  a  year,  or 
perhaps  a  little  longer,  but  there  are  some  whose  term  of  life 
extends  to  several  or  many  years. 

Trematodes  are  but  rarely  found  encysted  in  the  higher  verte- 
brate animals  ;  the  condition,  however,  is  more  frequent  in  amphi- 
bians, and  especially  in  fishes,  as  well  as  in  numerous  invertebrate 
animals. 

Classification  of  the  Trematodes  of  Man. 

The  following  classification,  partly  artificial,  partly  natural^ 
embraces  only  the  flukes  found  in  man  : — 

Order.     Digenea,  v.  Beneden,  1858. 

Anterior  sucker  single  and  median,  present.  Eggs  few.  The  (specialized) 
terminal  portion  of  the  uterus  serves  as  a  vagina.  Development  indirect,  i.e.^  an 
intermediate  host  is  required. 

Sub-order.     Prostomata,  Odhner,  1905. 

Mouth  surrounded  by  the  anterior  sucker. 

Group.     Amphistomata,   Rudolphi,  1801,  ep.,  Nitzsch,   1819. 

Gut  forked,  two  suckers,  the  posterior  sucker  (acetabulum)  terminal  or  ventro- 
terminal  behind  the  genitalia,  or  at  most  embraced  by  the  vitellaria.  Skin  with  no 
spines.     Excretory  bladder  a  simple  sac  opening  dorsally  near  hind  end.     Testes  in 


TREMATODES  :    CLASSIFICATION  23 1 

front  of  ovary.     Genital  pore,  median  in  anterior  third  of  body.    Thick  flukes,  ahnost 
circular  in  cross  section. 


Family.     Paramphistomidae,  Fischoeder,   1901. 

Amphistomata  :   Body  not  divided  into  a  conical  anterior  portion  and  disc-like 
caudal  portion.     Ventral  pouch  absent. 


Sub-family.     Paramphistominae,  Fisch.,  1901. 
ParamphistomidiE  :  Oral  sucker  without  evaginations.     Not  in  man. 

Sub-family.     Cladorchiinae,  Fisch.,   1901. 

Paramphistomidae :  Oral  sucker  with  evaginations ;  testes,  two,  deeply  cleft 
(fig-  I37)'     Genera:    IVatsonius,  Cladorckts,  tic. 

Family.     Gastrodlsciidae,  Stiles  and  Goldberger,  1910. 

Amphistomata  :  With  body  divided  into  a  conical  cephalic  and  disc-like  caudal 
portion  (fig.  138).  Posterior  sucker  ventro-terminal.  Oral  sucker  with  evaginations. 
Genera  :    Gastrodiscus  and  Homalogaster. 

Group.     Distomata,   Retzius,  1782. 

Gut  forked,  two  suckers,  the  posterior  sucker  (acetabulum)  ventral.  It  is 
always  separated  from  the  hind  end  by  at  least  a  part  of  the  genitalia. 

Family.     Fasciolidae,  Railliet,   1895. 

Large  flat  forms,  genital  pore  in  front  of  ventral  sucker,  the  latter  powerful. 
Vitellarias  of  numerous  follicles,  united  by  branching  vitellarian  ducts,  at  the  sides  of 
the  body  meeting  posteriorly  and  extending  ventrally  and  dorsally.  Cirrus  and 
vagina  without  spines.  No  crown  of  strong  spines  around  sucker.  Testes  much 
branched.  Uterus  not  well  developed.  Excretory  bladder  much  branched.  Eggs 
large. 

Sub-family.     Fasciolinae,  Odhner,  1910. 

Large  or  median  forms,  gut  much  branched.  Body  has  a  shoulder  separating 
head  from  body.  Receptaculum  seminis  absent.  Ovary  branched,  ventral  sucker 
in  anterior  part  of  body.     Genus :  Fasciola. 


Sub-family.     Fasciolopsinae,  Odhner,  1910. 

Shoulder  absent.  Receptaculum  seminis  present.  Ovary  branched,  gut  takes 
a  zig-zag  course  with  kinks  on  it,  ventral  sucker  in  anterior  part  of  body. 
Genus  :  Fasciolopsis. 


2^.2  THE   ANIMAL    PARASITES    OF    MAN 


Family.     Opisthorchiidae,  Braiin,  1901,  emend,  auctor. 

Ovary  in  front  of  testes.  Small  to  medium  flukes,  very  transparent,  tapering 
anteriorly.  Vitellaria  moderately  developed  not  extending  in  front  of  sucker. 
Cirrus  absent.  Seminal  vesicle  a  twisted  tube  free  in  parenchyma.  Testes  near 
hind  end  one  behind  the  other,  lobed  or  branched,  but  not  dendritically.  Excretory 
bladder  Y-shaped,  the  two  limbs  short,  the  stem  S-shaped  passing  between  the 
testes.  Receptaculum  seminis  well  developed.  Laurer's  canal  present.  Uterine 
coils  transverse,  numerous.     Eggs  small. 


Sub-family.     Opisthorchiinae,  Looss,  1899,  emend,  auctor. 

OpisthorchiidcE  in  which  the  excretory  pore  is  terminal.  Excretory  bladder  long, 
dorsal  to  testes.  Uterine  coils  not  overlapping  gut  forks.  Genera  :  Opisthorchis, 
Paropisthorchis,  Clonorchis,  Amphimerus,  etc. 


Sub-family.     Metorchiinae,  Liihe,  1909. 

Opisthorchiidcr  in  which  the  excretory  pore  is  ventral.  Excretory  bladder  short, 
ventral  to  testes.  Uterine  coils  partly  overlapping  gut  forks  and  extend  anteriorly 
beyond  the  sucker.   Vitellaria  compressed  on  the  sides  of  the  body.    Genus  :  Metorchis. 


Family.     Dicrocoeliidae,  Odhner,   1910. 

Ovary  behind  testes.  Testes  behind  the  ventral  sucker,  between  it  and  the  ovary. 
Body  thin  and  transparent.  Cirrus  sac  encloses  the  pars  prostatica  and  seminal 
vesicle.  Skin  smooth.  Gut  forks  do  not  reach  posterior  end.  Receptaculum  seminis 
and  Laurer's  canal  present.  Vitellaria,  moderate,  lateral  in  mid-body  slightly  over- 
lapping the  gut.  Uterus  with  an  ascending  and  descending  branch  and  numerous 
transverse  coils  extending  to  hind  end.  Eggs  dark  brown,  25  ju  to  60  m.  Excretory 
bladder  tubular  in  posterior  third  or  half  of  body.  Parasitic  in  bile-ducts  of  mammals 
and  birds.    Genus  :    Dicrocoeliuin, 


Family.     Heterophyiidae,  Odhner,  1914. 

Ovary  in  front  of  testes.  Genital  pore  behind  or  on  a  level  with  ventral  sucker. 
Genital  pore  surrounded  by  a  pseudo-sucker  {i.e.,  its  muscle  is  not  sharply  separated 
from  but  blends  with  the  body  muscles).  Cirrus  sac  absent,  consequently  vesicula 
seminalis  and  pars  prostatica  lie  free.  Vagina  and  ejaculatory  duct  unite  into  a 
common  duct  before  opening.  Small  and  very  small  forms.  Body  covered  with 
scales.     Genera  :  Heterophyes,  Meiagonimus^  etc. 


Family.     Troglotremidae,  Odhner,  1914. 

More  or  less  flattened  Distomes  of  compact  form,  2  to  13  mm.  long.  Ventral 
surface  flat  or  somewhat  hollowed,  dorsal  surface  arched.  Skin  completely  covered 
with  pointed  spines.      Musculature  weakly  developed  also  in  the  suckers  in  those 


TREMATODES  :     CLASSIFICATION 


233 


forms  that  inhabit  cysts.  Gut  with  pharynx  and  a  not  very  long  oesophagus  and 
caeca,  which  end  more  or  less  shortly  before  the  hind  end.  Excretory  bladder 
Y-shaped  or  tubular.  Pars  prostatica  and  seminal  vesicle  always  distinct.  Testes 
elongated,  symmetrically  placed  in  or  behind  the  middle  of  the  body.  Ovary  directly 
in  front  of  the  testes,  right-sided,  generally  much  lobed.  Receptaculum  seminis 
and  Laurer's  canal  present.  Vitellaria  generally  well  developed,  exclusively  or  for 
the  most  part  confined  to  the  dorsal  surface^  leaving  only  a  median  band  unoccupied. 
Uterus  either  very  long,  coiling  here  and  there,  or  shorter  and  more  convoluted. 
Eggs  in  first  case  small  17  ju  to  25  ^u,  in  the  second  much  larger  63  a*  to  85  /x  or 
even  120 /*(?)  long.  Parasitic  in  carnivora  or  birds,  generally  occurring  in  pairs 
in  cyst-like  cavities.     Genera  :  Paragonhnus^  Pholeter^  Collyriclum,  Troglotreina. 


Family.     Echinostomidae,  Looss,  1902. 

More  or  less  elongated  flukes^  small  or  very  large,  much  flattened  anteriorly, 
less  so  posteriorly,  or  even  round.  Suckers  near  one  another,  the  anterior  small  and 
weak,  the  posterior  large  and  powerful  directed  obliquely  backwards.  Stcrrounding 
the  oral  sucker  dorsally  and  laterally  but  7tot  venirally  is  a  fold  or  "  collar ''  bearing 
a  row  or  rows  of  pointed  spi?tes  which  are  continued  round  laterally  o?i  to  the  ventral 
corners,  the  number  being  constant  for  each  species,  the  corner  spities  large  or 
specialized,  skin  anteriorly  scaled  or  spiny.  Alirne7ttary  canal  consists  of  a  pharynx, 
epithelial  '''•  pseudo-ossophagus  "  and  gut  cceca  reaching  to  posterior  end.  Testes  behind 
one  a?tother  in  hind  body.  Ovary  on  right  side  or  median  directly  in  front  of  the 
testes.  Vitellaria  lateral,  usually  extending  to  the  hind  end  and  not  beyofid  the 
ve?ttral  sucker  anteriorly.  Genital  pore  fust  in  front  of  ventral  sucker.  Uterus  i?i 
transverse  loops.  Genital  sinus  absent  or  present.  Receptaculum  seminis  and 
Laurer's  canal  prese?it.  Eggs  thin  shelled  and  large,  bright  yellow,  6$  /x  to  120  /j.  long. 
Excretory  bladder  ^ -shaped.     Parasitic  in  gut  of  vertebrates,  especially  birds. 


Sub-family.     Echinostominae,  Looss,  1899. 

Cirrus  sac  usually  reaching  to  centre  of  ventral  sucker,  but  not  beyond.  Cirrus 
long,  usually  without  spines,  coiled  when  retracted.  Sejninal  vesicle  tubular,  twisted. 
On  the  head  a  ventral  uniting  ridge  between  the  angles  of  the  collar.  Dorsal  circlet 
of  spines,  siftgle  or  double,  not  interrupted  unless  the  collar  itself  is  dorsally  divided. 
Genera :  Echinostoma,  etc. 


Sub-family.     Himasthllnae,  Odhner,  1910. 

Cirrus  sac  reaching  far  beyond  ventral  sucker.  Cirrus  armed  with  strong 
rose-thorn-shaped  hooks.  Vesicula  seminalis  tubular  not  coiled.  Cervical  collar 
not  continued  across  ventral  aspect.  Spines  on  collar  in  one  row.  Body  armed 
with  fine  needle-shaped  spines. 


Family.     Schistosomidae,  Looss,  1899. 

Sexes  separate.  Genital  pore  behind  the  ventral  sucker.  Ventral  sucker 
elevated  above  the  surface.  Pharynx  absent.  Gut  forks  reunite  to  form  a  single 
stem.  In  $  four  or  more  testicular  follicles.  In  ?  a  single  ovary,  just  in  front 
of  the  union  of  the  gut  forks.     Vitellaria  on  either  side  of  the  united  gut  stem. 


234 


THE   ANIMAL   PARASITES   OF   MAN 


The  Trematodes  Observed  in  Man. 
Family.     Paramphistomldae,  Stiles  and  Goldberger,  emend.  1910. 
Sub-family.     Cladorchiinae,   Fisch.,  1901. 
Genus.     Watsonius,  Stiles  and  Goldberger,  1910. 

CladorchmcE.—^ody  pyriform.  Ventral  pouch  absent.  Acetabulum  ventral  or 
(?)  ventro-subterminal,  very  large,  margins  projecting,  aperture  small.  Genital 
pore  in  front  of  bifurcation  of  gut,  not  surrounded  by  a  sucker  ;  ductus  herma- 
phroditicus  apparently  absent.  Excretory  pore  at  posterior  end  of  excretory  vesicle, 
behind  Laurer's  canal.  Oral  sucker  with  a  pair  of  irregularly  globular  suctorial 
pouches  ;  oesophagus  thickened  distally  ;  caeca  long,  not  wavy  ;  end  in  acetabular 
region. 

Male  Or£-a;is.— Testes  two  lobed,  smaller  than  acetabulum  ;  longitudinally, 
nearly  or  quite  coinciding  ;  transversely  they  abut  or  slightly  overlap  ;  preovarial 
in  equatorial  and  caudal  thirds.  Pars  musculosa  not  largely  developed;  cirrus 
pouch  absent. 

Female  Organs. — Ovary  and  shell  gland  post-testicular.  Vitellaria  extend  from 
gut  fork  to  slightly  beyond  gut  ending  ;  uterus  interccecal,  partly  post-testicular. 
Laurer's  canal  in  front  of  excretory  vesicle. 

Type  Species. —  Watso?tius  watsoni,  Conyngham,  1904. 


Watsonius  watsoni,  Stiles  and  Goldberger,  1910. 

Syn. :   Ainphistojuuni  -£v^/J•<?«^,  Conyngham,  1904;  Cladorchis  watsoni^  Shipley,  1905. 

Body,  8  to  10  mm.  long,  by  4  to  5  mm.  broad,  by  4  mm.  thick ;  tapers 
anteriorly  to  2*5  mm.  Caudal  extremity  bluntly  rounded,  venter 
surrounded  by  an  elevated  ridge,  surface  with 
transverse  ridges  best  defined  ventrally.  Genital 
pore  median  about  one-quarter  of  body  length 
from  anterior  end  at  level  of  suctorial  pouches. 
Acetabulum  i  mm.  in  diameter,  margin  project- 
ing, aperture  small.  Mouth  in  a  groove  with 
digitate  papillae.  Oral  sucker  very  large,  one-fifth 
I  j      of    length    of    body,   witli    a   pair   of  irregularly 

I  I      globular  pouches.     QEsophagus  somewhat  longer 

than  sucker.  Excretory  pore  at  the  level  of 
the  acetabular  aperture.  The  vesicle  extends 
from  the  plane  of  the  transverse  vitelline  ducts 
to  centre  of  acetabulum. 

Male  Organs. — Testes  deeply  notched  adjoining 

one   another.      Vesicula   seminalis    much    coiled 

and    dilated,  pars  musculosa  not  coiled.     Pars  prostatica  (?)  dilated, 

ejaculatory   duct    long   and    narrow,  opening   on   a  papilla ;   genital 

atrium  papillated. 


Fig.  136. —  Walsonius 
watsoni:  ventral  view. 
4/1.     (After  Shipley.) 


WATSONIUS   WATSONI 


=  35 


Female  Organs. — Ovary  dorso-posterior  of  posterior  testis.  Shell 
gland  dorsal  to  ovary.  Vitellaria  ventral  and  lateral  to  gut  caeca 
extending  from  gut  fork  to  equator  of  acetabulum.  Uterus  dorsal 
to  testes,  ductus  hermaphroditicus  absent.  Laurer's  canal  opens  in 
dorso-median  line  slightly  behind  anterior  border  of  sucker. 


Fig.  iTfj.—  IVatsonitts  watsoni :  ventral  projection  composed 
from  a  series  of  transverse  sections,  o.s.,  oral  sucker  ;  s.p.,  suc- 
torial pouch;  ga.^  genital  atrium;  d.e.,  ejaculatory  duct; 
(?5.,  oesophagus ;  e.g.,  oesophageal  ganglion  ;  p.p.,  pars  prostatica  ; 
p.vi.,  pars  musculosa  ;  i,,  gut;  w/.,  uterus;  v.e.,  vas  efferens; 
v.e.s.,  left  vas  efferens  ;  v.e.d.,  right  vasefiferens  ;  v.g.,  vitellarium  ; 
/.,  testes  ;  ov.,  ovary  ;  s.g.,  shell  gland  ;  t.vd.,  transverse  vitelline 
duct.     (After  Stiles  and  Goldberger.) 


Eggs. — 123  yLt  to  133  yu-  long  by  75  /jl  to  So  fi  broad. 

Habitat. — Jejunum  and  duodenum  of  man,  German  West  Africa. 
The  parasite  has  only  been  found  once  in  man.  The  patient,  a  negro 
from  German  West  Africa,  died  at  Zola,  Northern  Nigeria.  The 
symptoms  were  persistent  watery  diarrhoea  without  blood  or  mucus. 
The  parasites  were  also  passed  in  the  stools.  It  occurs  also  in 
monkeys. 


236  THE   ANIMAL   PARASITES   OF   MAN 

Family.     Gastrodisciidae. 
Genus.     Gastrodiscus,  Lkt.,  1877. 

Acetabulum  small,  caudal  and  ventral  margin  raised,  aperture  relatively  large. 
Genital  pore  without  sucker.  Excretory  pore  post-vesicular,  posterior  to  opening 
of  Laurer's  canal.  (Esophagus  with  muscular  thickening  ;  caeca  not  wavy,  long, 
end  post-equatorial  and  post-testicular. 

Male  Genitalia. — Testes  two,  branched  pre-ovarial. 

Female  genitalia. — Ovary  and  shell  gland  post-testicular.  Vitellaria  extracsecal  ; 
uterus  intercaecal  ;  Laurer's  canal  entirely  prevesicular. 

Type. — Gastrodiscus  cBgyptiacus^  Cobbold,  1876. 


Gastrodiscus  hominis,  Lewis  and  McConnell,  1876.^ 
Syn.  :   Amphistoimim  ho^ninis^  Lew.  and  McConn. 

Body,  reddish   in   the  fresh,  5    to   8  mm.  long;    posteriorly,  3  to 

4  mm.  broad.     The  disc  has  incurved  edges  which  are  interrupted  in 

front  where  it  joins  the  anterior  cylindrical  portion  and  posteriorly 

behind  the  ventral   sucker.      The  disc  itself  and  ventral  surface  are 

covered  with   a   number  of   (microscopic)  papillae. 

Pharynx  provided  with  two  diverticula  or  pouches. 

The  bifurcation  of   the  gut   lies  sometimes  above, 

sometimes  below  the  level  of  the  genital  pore.     The 

gut  caeca  end  about  the  level  of  the  centre  of  the 

Ik         Jl         acetabulum. 

1^1        ^1  Genital  Pore. — About  the  middle  of  the  conical 

^^        anterior  portion.  (It  appears  to  be  surrounded  by  a 

dJl^i\^\'7mTni's      n^uscular  suckcr.)     Leiper  (1913)  describes  the  ducts 

Slightly     magnified,     as  discharging  at  the  tip  of  a  large  fleshy  papilla,  the 

(After  Lerckart.)         surface  of  which  bears  cuticular  bosses. 

Testes  much  lobed,  the  anterior  is  smaller  than  the  posterior  and 
lies  at  about  the  level  where  the  anterior  conical  portion  joins  the 
disc.  The  posterior  testis  just  in  front  of  the  anterior  margin  of  the 
acetabulum  separated  from  it  by  the  ovary.  The  ovary,  somewhat 
oval  in  shape  or  slightly  constricted  in  the  middle,  lies  slightly  to  the 
right  of  the  median  line.  Dorsal  to  it  lies  the  well-developed  shell 
gland,  Laurer's  canal  opening  in  front  of  the  excretory  bladder. 
The  excretory  bladder  is  a  long  sac  with  its  openmg  at  its  posterior 
extremity   about   the   level   of   the  middle  of   the  acetabulum.     The 


*  Leiper  places  this  species  in  a  new  genus  Gastrodiscoides.  Genus  Gastrodiscoides,  Leiper, 
1913,  distinguished  from  Gastrodiscus  by  :  (i)  large  genital  cone  ;  (2)  position  of  genital  orifice  ; 
{3)  disc  without  papillae  ;  {4)  testes  one  behind  the  other. 


FASCIOLID^i 


237 


vitellaria  are  restricted  in  extent.  They  do  not  extend  forward 
beyond  the  anterior  border  of  the  posterior  testis.  They  are  best 
developed  in  the  area  between  the  acetabulum  and  the  termination 
of  the  gut  caeca. 

The  eggs  are  oval  and  measure  150  /z-  in  length  by  72  /-t  in 
breadth. 

Habitat. — Caecum  and  large  intestine  of  man.  Also  in  the  pig 
(5  per  cent.)  in  An  nam. 

Distribution. — This  parasite  has  been  recorded  from  Assam  (not 
uncommon),  British  Guiana  (Indian  immigrants),  and  Cochin  China. 

Gastrodiscus  a^gyptiaciis,  Cobbold,  1876,  and  G.  secundus,  Looss, 
1907,  occur  in  the  horse;  G.  minor,  Leiper,  1913,  in  the  pig  in 
Nigeria  and  Uganda. 


Family.     Fasciolidae,  Raill.,  1895. 

Sub-family.     Fasciolinae,  Odhner,  1910. 

Genus.     Fasciola,  L.,  1758. 

The  ventral  sucker  is  situated  at  the  level  of  the  junction  of  the  cone  with  the 
body,  viz.,  at  the  level  of  the  "shoulder,"  and  is  large  and  powerful.  The  cuticle 
is  covered  with  strong  spines  ;  the  gut  caeca  run  in  the  mid-line  to  the  hind  end, 
and  are  provided  with  numerous  long  lateral  and  fewer  and  shorter  median  branches. 
The  ovary  lies  on  one  side  in  front  of  the  transverse  vitelline  duct ;  the  testes 
lie  obliquely  one  behind  the  other.  The  uterus,  in  the  shape  of  a  rosette,  lies 
in  front  of  the  genitalia.  Laurer's  canal  is  present  ;  the  vesicula  seminalis  lies 
in  the  cirrus  pouch  ;  the  ova  are  large,  not  very  numerous,  and  only  develop  after 
they  have  been  deposited.     Parasites  of  the  biliary  ducts  of  herbivorous  animals. 


Fasciola  hepatica,  L.,  1758. 

Syn.  :   Distommn  hepaticmn,  Retz.,  1786  ;  Fasciola  humana,  Gmel.,  1789  ; 
Distomuvi  cavicB,  Sons.,  1890  :  Cladocoeliiaii  hepaticum,  Stoss.,  1892. 

Length  20  to  30  mm.,  breadth  8  to  13  mm.,  cephalic  cone  4  to  5  mm. 
in  length  and  sharply  differentiated  from  the  body  by  a  shoulder  on 
each  side.  Spines  in  alternating  transverse  rows  and  extending  on 
the  ventral  surface  to  the  posterior  border  of  the  testes,  and  on  the 
dorsal  surface  not  quite  so  far.  The  spines  are  smaller  on  the  cephalic 
cone  than  on  the  posterior  part  of  the  body,  where  they  are  discernible 
with  the  naked  eye.  The  suckers  are  hemispherical,  and  near  each 
other ;  the  oral  sucker  is  about  i  mm.  and  the  ventral  sucker  about 
1*6  mm.  in  diameter.     The  pharynx,  which  includes  almost  the  entire 


-238 


THE   ANIMAL   PARASITES   OF   MAN 


oesophagus,  measures  07  mm.  in  length  and  0*4  mm.  in  breadth. 
The  intestine  bifurcates  at  the  Umit  of  the  cephahc  cone  and  the 
branches  are  even  here  furnished  with  diverticula  directed  outwardly. 
The  ovary  is  ramified  and  situated  in  front  of  the  transverse  vitelline 
duct,  usually  on  the  right  side;  the  shell  gland  lies  near  the  ovary  in 
the  median  line ;  posterior  to  the  transverse  vitelline  ducts  are  the 
greatly  ramified  testes,  which  occupy  the  greater  portion  of  the 
posterior  part  of  the  body,  with  the  exception 
of  the  lateral  and  posterior  border  ;  the  long 
vasa  efferentia  only  unite  as  they  enter  the 
cirrus  pouch.  The  vitellaria  occupy  the  sides 
of  the  posterior  part  of  the  body,  commencing 
at  the  level  of  the  ventral  sucker  and  uniting 
behind  the  testes.  The  ova  are  yellowi^-brown, 
oval,  operculated,  130  ^  to  145  //<  in  length,  70  {x 
to  90  /A  in  breadth  (average  size  132  /a  by  70  fi). 
The  Liver  Fluke  inhabits  the  bile-ducts  of 
numerous  herbivorous  mammals  (sheep,  ox, 
goat,  horse,  ass,  rabbit,^  guinea-pig,  squirrel, 
beaver,  deer,  roe,  antelope,  camel,  kangaroo, 
and  others),  and  is  distributed  over  the  whole 
of  Europe,  though  not  to  an  equal  extent.  It 
is  further  known  in  North  Africa,  in  North  and 
South  America,  as  well  as  in  Australia ;  it  is 
also  found  in  Asia,  as  it  has  been  reported 
from  Japan,  China,  and  Tonkin  (Gaide,  two 
cases  in  man).  In  some  districts  of  Germany 
it  is  very  frequent,  and  the  slaughter-house 
statistics  of  various  places  show  that  it  is  of 
daily  occurrence.  Fasciola  magna  occurs  in 
herbivora  in  America. 

The  liver  fluke,  however,  is  by  no  means  a 
harmless  parasite,  for  it  produces  in  domestic 
animals,  more  especially  in  sheep,  a  disease  of  the  liver  that  appears 
epidemically  in  certain  years  and  districts,  and  commits  great  ravages 
amongst  the  flocks. 

[The  following  records  show  the  enormous  loss  caused  in  sheep 
by  this  parasite.  In  181 2,  in  the  Midi,  principally  in  the  Departments 
of  the  Rhone,  Herault,  and  Gard,  the  disease  was  rampant;  300,000 
sheep  perished  in  the  Aries  territory,  and  90,000  in  the  Arrondissements 
of  Nimes  and  Montpellier.      In   1829  and   1830,  in  the  Department 


Fig.  IT,().— Fasciola  hepa- 
iica,  L.  From  a  specimen 
that  is  not  yet  mature, 
showing  the  gut  and  iis 
branches.     5/1. 


'  [There  does  not  seem  to  be  any  direct  evidence  of  either  rabbits  or  hares  normally 
being  invaded  by  this  fluke. — F.  V.  T.] 


FASCIOLA    HEPATICA 


239 


of  the  Meuse  and  near  localities,  not  only  sheep  but  oxen  died  in 
enormous  numbers;  for  instance,  in  the  Arrondissement  of  Verdun 
out  of  50,000  sheep  20,000  died,  and  out  of  20,000  cattle  2,200  died. 
In  England,  in  1830,  2,000,000  sheep  were  carried  off;  whilst  in  1862 
60  per  cent,  of  the  sheep  died  in  Ireland;  and  in  1879  over  300,000 
were  lost  in  England ;  whilst  as  late 
as  189 1  one  owner  in  the  same 
country  lost  over  10,000  sheep  {Live 
Stock  Journal,  October  30,  189 1). — 
F.  V.  T.] 

The  disease  usually  commences 
towards  the  end  of  summer  with  an 
enlargement  of  the  liver,  induced  by 
the  invasion  of  numerous  young 
flukes;  in  the  autumn  and  winter  the 
animals  suffer  from  the  consequences 
of  disordered  biliary  secretion ;  they 
become  feverish,  emaciated,  and 
anaemic,  and  lose  their  appetite. 
In  consequence  of  the  consecutive 
atrophy    of    the    liver,    oedema    and 


Ov. 


E.d. 


Fig.  140. — Fasciola  hepatica.  M.,  mouth; 
Ut.^  uterine  rosette  ;  Tr.c,  transverse  vitelline 
ducts  uniting  to  form  a  vitelline  receptacle  in 
the  mid-line  ;  E.d.,  longitudinal  vitelline  ducts  ; 
F.i-.,  vitellaria.  The  clear  space  in  the  centre 
represents  the  position  of  the  ramifying  testes 
and  part  of  the  gut.  Natural  size.  (Mull,  fluid, 
alcohol,  creosote,  Canada  balsam.) 


^FiG.  141. — Fasciola  hepatica^  L.  /., 
intestine;  Fj.,  vitellaria;  Ov.,  ovary; 
O.,  oral  aperture;  Ut.^  uterus;  S.,  ven- 
tral sucker  ;  7".,  testes.  In  front  of  the 
testes  are  seen  the  transverse  vitelline 
ducts  uniting  to  form  the  pyriform  vitel- 
line receptacle.  Immediately  in  front  of 
this  the  spherical  shell  gland.  The  two 
vasa  efferentia  can  also  be  seen  running 
up  in  the  mid-line.  The  branches  of  the 
gut  are  only  shown  in  the  cephalic  cone. 
(After  Claus.) 


ascites  set  in,  and  many  animals  succumb  to  this  '^  liver  rot."  On 
exammation  the  liver  is  found  to  be  shrunken,  the  bile-ducts  are 
enormously  dilated  and  in  parts  saccular  and  full  of  flukes.  Should 
the  animals  survive  this  stage,  spontaneous  recovery  ensues  in 
consequence    of   the   flukes   commencing   to   leave   the   liver  in   the 


240 


THE   ANIMAL   PARASITES   OF   MAN 


spring,  but  the  liver  remains  changed  and  its  sale  is  prohibited^  when 
the  changes  are  extensive.^ 

[The  following  stages  may  be  noticed  in  sheep  suffering  from 
fascioliasis.  Gerlach  recognized  four  stages,  based  on  the  varied 
relations  that  the  flukes  contract  with  the  liver  of  their  host.  These 
periods  are  sometimes  very  marked,  but  at  others,  owing  to  subsequent 
infections,  the  features  become  merged  and  so  obliterated.  But 
when  a  single  infestation  occurs  they  are  very  marked. 

[The  first  period  is  called  the 

PERIOD    OF     IMMIGRATION.       This 

occurs  at  the  fall  of  the  year  and 
generally  passes  unperceived^  as 
the  young  flukes  do  little  harm 
to  the  liver.  It  varies  from  four 
to  thirteen  weeks.  Gerlach  has 
remarked  upon  cases  of  death 
from  apoplexy  at  this  period. 

[The  second  period  is  the 
PERIOD  OF  ANi4:MiA.  This  occurs 
in  November  and  December. 
The  sheep  at  fust  fatten  rapidly, 
but  later  the  mucous  membranes 
become    pale  and  of  a  yellowish 


Fig.  142. — Fasciolahepatical  egg  from  liver 
of  sheep.  0,  operculum,  e,  segmenting  ovum. 
The  rest  of  the  space  is  occupied  by  yolk  cells, 
the  granules  in  three  only  being  shown,  x  68c. 
(After  Thomas.) 


Fig.  143. — Limvaus  trnncatulus,  Mlill., 
the  intermediate  host  of  Fasciola  hepatica. 
a.,  natural  size ;  d.,  magnified.  (From 
Leuckart.) 


hue,  and  the  sheep  become  sluggish  and  cease  to  feed.     The  faeces 
are  normal,  but  may  contain  fluke  ova. 

[The  third  period  is  the  PERIOD  OF  wasting.  This  corresponds 
with    the    beginning    of    January — about    three    months    after    the 

'  [This  is  not  the  case  in  Great  Britain  ;  fluky  sheep  are  sent  to  market,  there  being  no 
danger  to  man  from  eating  the  flesh. — F.  V.  T.] 

2  As  an  example,  this  occurred  in  Berlin  in  the  case  of  19,034  oxen,  15,542  sheep,  1,704 
pigs,  and  160  calves  in  the  period  of  1883-1893  ;  during  which  time  719,157  oxen,  1,519,003 
sheep,  2,258,110  pigs,  and  567,964  calves  were  slaughtered.  As  a  matter  of  fact,  however, 
the  number  of  infected  beasts  was  really  larger. 


FASCIOLA   HEPATICA  24T 


entry  of  the  larvae.  Emaciation  now  becomes  very  marked,  the 
skin  and  mucous  membranes  blanched,  temperature  variable  and 
marked  by  an  irregular  curve ;  respiration  laboured  and  quick ; 
appetite  regular;  abortion  frequently  occurs  in  pregnant  ewes; 
pressure  on  the  back  causes  the  animals  to  fall ;  local  oedemas 
occur,  the  most  perceptible  in  the  submaxillary  space,  extending 
below  the  larynx  and  over  the  cheeks  and  parotids  (called 
**  bourse,"  *' boule  "  in  France;  "watery  poke"  or  '^  cockered"  in 
England).  Death  usually  occurs  at  this  period,  but  a  fourth 
stage  may  occur. 

[The  fourth  period  is  the  PERIOD  OF  migration  of  the 
FLUKES.  This  is  a  period  of  convalescence  and  recovery,  generally 
in  May  and  June.— F.  V.  T.] 

Oxen  suffer  less  in  general,  but  even  in  these  animals  ''  stray " 
hepatic  flukes  are  occasionally  found  in  the  lungs,  enclosed  in  thick- 
walled  cysts. 

Pathological  Anatomy. — The  bile-ducts  are  conspicuous  on  the 
surface  of  the  liver.  They  are  thickened  and  much  dilated  and  in 
parts  saccular,  and  considerable  atrophy  of  the  liver  cells  accom- 
panies the  condition.  Histologically  there  is  immense  proliferation 
of  the  epithelium  of  the  bile-ducts  leading  to  "adenomata." 

The  LIFE -HISTORY  of  the  liver  fluke  was  discovered  by 
R.  Leuckart  and  P.  Thomas.  According  to  these  investigators  the 
elongated  miracidium  (fig.  131,  a)  ciliated  all  over  develops  from  the 
eggs  a  few  weeks  after  the  latter  (fig.  142)  have  reached  the  water,  and 
after  it  has  become  free  the  embryo  penetrates  and  becomes  a  sporo- 
cyst  (fig.  131,  b)  in  a  water-snail  {Limnceus  trmtcatulus,  Miill.  = 
L.  niinutus,  Drap.)  that  is  common  in  fresh  water,  and  can  live  m  the 
smallest  collection  of  water  as  well  as  in  fields  that  have  been  flooded. 
The  sporocyst  first  of  all  produces  rediae,  which  remain  in  the  same 
host  (and  under  certain  circumstances,  e.g.  in  summer,  these  develop  a 
second  generation  of  rediae),  and  these  finally  form  cercariae  (fig.  134). 
The  latter  become  encysted  on  blades  of  grass  and  are  taken  up  by 
the  respective  hosts  with  their  food;  this  takes  place  towards  the 
end  of  summer,  while  the  sheep  feeding  on  the  pasture  land  in  the 
spring  spread  the  eggs  of  the  fluke,  and  sometimes  the  fluke  itself, 
by  passing  them  with  their  faeces. 

In  districts  where  Limnceus  truncatultis  is  absent,  analogous 
species  act  as  the  intermediary  hosts,  of  which  one  example 
according  to  Lutz  is  Limnceus  oahuensis  in  the  Sandwich  Islands. 

[The  host  in  Europe  is  Limnceus  truncatultis.  This  snail 
extends  from  Siberia  to  Sicily  and  Algeria,  and  according  to 
•Captain  Hutton  is  a  native  of  Afghanistan.  It  also  occurs  in 
16 


242  THE   ANIMAL   PARASITES   OF   MAN 

Thibet,  Amoor,  Morocco,  Tunis,  Canary  Islands  and  the  Faroe 
Islands.  It  deposits  its  eggs  or  spawn  upon  the  mud  around  ponds, 
ditches  and  streams.  The  eggs  are  laid  in  batches  of  thirty  to  a 
hundred,  each  snail  laying  as  many  as  1,500  eggs  ;  they  are  united 
into  strips  of  a  gelatinous  substance.  In  about  two  weeks  young 
snails  appear.  It  is  amphibious,  being  more  frequently  met  with  out 
of  the  water  than  in  it.  It  occurs  in  elevated  spots  as  well  as  in 
low-lying  districts.  Moquin-Tandon  found  it  at  4,000  feet  in  the 
Pyrenees.  In  the  allied  species,  L.  peregra,  the  fluke  will  develop 
up  to  a  certain  stage,  but  never  completes  all  its  varied  phases. 

[In  South  America  the  host  is  probably  Limnceus  viator.  Orb., 
and  in  North  America  Limnceus  ImmiliSy  Say. — F.  V.  T.] 

In  human  beings  as  well  as  in  some  of  the  mammals  quoted 
above,  the  liver  fluke  is  only  a  casual  parasite,  and  hitherto  only 
twenty-eight  cases  have  been  observed  in  man  ;  the  infection  was 
mostly  a  mild  one  and  there  were  no  symptoms,  or  only  very 
trifling  ones ;  a  few  isolated  cases  were  only  discovered  post  mortem. 
Occasionally,  however,  even  when  the  infection  was  inconsiderable, 
severe  symptom.s  were  set  up,  which  in  isolated  cases  led  to  death. 
The  symptoms  (enlargement  and  painfulness  of  the  liver,  icterus) 
merely  pointed  to  a  disease  of  the  liver. 

Diagnosis  can  only  be  established  by  finding  eggs  in  the  faeces. 
Care  should  be  taken  not  to  confuse  them  with  those  of  Dibothrio- 
cephalns  latus. 

Halzoun. 

In  North  Lebanon,  the  liver  fluke  is,  according  to  A.  Khouri, 
a  frequent  parasite  of  man,  not  in  the  liver,  however,  but  in  the 
pharynx.  The  occurrence  in  this  unusual  site 
is  effected  by  the  eating  of  raw  infected  livers, 
especially  those  of  goats  (Capra  hirciis).  The 
flukes  thus  taken  in  do  not  all  reach  the 
stomach,  where  they  would  be  soon  killed,  but 
some  of  them  attach  themselves  to  the  pharyn- 
geal mucosa  and  to  the  adjoining  parts,  and 
there  cause  inflammation  and  swelling,  which 
lead  to  dyspnoea,  dysphagia,  dysphonia  and 
Fig.  144.— Young  Fas-      congestion    of    the    head,    sometimes   even  to 

ciola    hepaitca,   soon   after  ,.,,  '  ,  ,       ,, 

entry  into  the  liver.    The      stili   more    severe  symptoms,  and  even  death, 
intestinal  caeca  have  lateral      The  affection   termed   '' Halzoun  "  lasts    some 

diverticula.         Magnified.         , 

(From  Leuckart.)  tiours    or    several    days,    and    after    vomitmg 

recovery  sets  in.     In  other  cases  man  becomes 

infected  in  the  usual  way   by  ingesting   cysts   attached   to  grass  or 

the    underside   of   leaves   of    plants    {e.g.,    Rumex    sp.),    where   they 


HALZOUN 


243 


are  overlooked  from 
their  scanty  size  (0*2  to 
o'3  mm.). 

As  the  liver  fluke 
feeds  on  blood  it  is 
possible  that  it  also 
reaches,  particularly 
when  young,  the  circu- 
latory system,  and  cases 
have  been  known  in 
which  it  has  been  carried 
by  the  blood  into  organs 
far  from  its  original  situ- 
ation. Such  cases  also 
have  been  repeatedly 
observed  in  men.  Pro- 
bably the  parasite  de- 
scribed by  Treutler, 
1793,  as  Hexathyridmm 
venaruntf  which  pro- 
truded from  the  rup- 
tured anterior  tibial  vein 
of  a  man,  was  a  young 
liver  fluke.  A  few  adult 
specimens  were  found 
by  Duval  in  the  portal 
and  other  veins  post 
mortem  at  Ren nes  (1842) 
in  a  man,  aged  49,  and 
a  similar  statement  is 
reported  by  Vital  from 
Constantine  (1874). 
Giesker,  in  1850,  found 
two  hepatic  flukes  in  a 
swelling  on  the  sole  of 
the  foot  of  a  woman. 
Penn  Harris  states  that 
he  observed  six  speci- 
mens in  Liverpool  in  a 
spontaneously  ruptured 
abscess  of  the  occiput 
of  a  two  months  old 
infant.  Another  case 
which,  like  the  previous 


Genital 


Vitellariutn 


Posterior . 
testis 


Uterus 


,    Ovary 


Anterior 
testist  I 


Excretory  pore 


Fig.  145. 


-Fasciola  gigantica. 
(After  Looss.) 


6^. 


24^  THE   ANIMAL   PARASITES   OF   MAN 

one,  is  reported  by  Lankester,^  relates  to  a  sailor  who  suffered  from 
an  abscess  behind  the  ear,  and  from  which  a  Hver  fluke  was  expelled. 
Finally,  Dionis  de  Carrieres  reports  the  case  of  a  man,  aged  35,  in 
whose  right  hypochondriac  region  a  tumour  the  size  of  a  pigeon's  egg 
had  formed,  and  from  which  a  young  liver  fluke  was  extracted. 

From  such  records  it  is  not  impossible  that  Distomum  oculi 
humani,  Ammon,  1833,  as  well  as  Monostomum  lentis,  v.  Nordm., 
1832,  may  have  been  very  young  hepatic  flukes  that  had  strayed. 
.Ammon  found  four  specimens  (length  0*5  to  i  mm.)  of  his  species 
(named  Distomum  ophthalmohium  by  Diesing  in  1850)  between  the 
opaque  lens  and  the  capsule  of  a  five  months  old  child  in  Dresden, 
and  von  Nordmann  discovered  his  Monoslomiim  lentis  to  the  number 
of  eight  specimens  (only  0*3  mm.  in  length)  in  the  opaque  lens  of 
an  old  woman.  Minute  white  bodies  which  Greef  found  in  the  cortex 
of  the  lens  of  a  fisherman,  aged  55,  removed  on  account  of  cataract, 
were  with  some  reserve  regarded  as  Trematode  larvae.  The  fact  that 
Ammon  found  that  the  intestinal  caeca  of  the  worm  discovered  by 
him  had  no  lateral  branches  does  not  negative  the  above  opinion,  as 
in  the  liver  fluke  the  intestinal  caeca  are  originally  unbranched,  and 
according  to  Lutz  they  only  develop  lateral  ramifications  later,  between 
the  twelfth  and  twent^^-second  day  of  infection  (fig.  144). 


Fasciola  gigantica,  Cobbold,  1856. 

Syn.  :  Disto?num  giganteufn,  Diesing,  1858;  Fasciola  gigantea^  Cobbold,  1858  ; 
Cladocoelium  giganteum^  Stoss.,   1892  ;  Fasciola  hepatica  var.  angusta^  Raill.,  1895  \ 

•rin]n  hd>'h/itirn  \jCkY     /vcrM'htinr/i     T  rvricc      iRnA 


Fasciola  hepatica  var.  cBgyptiaca^  Looss,  1 896. 


This  species  is  closely  allied  to  Fasciola  hepatica,  but  is  distin- 
guished by  its  elongated  body,  short  cephalic  cone,  almost  parallel 
sides,  larger  ventral  sucker,  which  is  also  closer  to  the  oral  sucker, 
and  by  its  larger  eggs.  Length  up  to  75  mm.,  width  up  to  12  mm. 
Oral  sucker  i  to  1*2  mm.,  ventral  sucker  up  to  17  mm.  in  diameter. 
Eggs  150  />fc  to  190  /A  long  by  75  //,  to  90  //,  broad. 

Habitat. — Bile-ducts  of  Camelopardalis  giraffaj  Bos  taunts,  Bos 
indicus,  Bos  bubalis^  Ovis  aries  and  Capra  hitcns. 

Distribution. — Africa. 

This  species  has  once  been  observed  in  man  by  Gouvea,  in  Rio  de 
Janeiro,  in  a  French  naval  officer  who  became  ill  with  fever,  cough 
and    slight    blood-spitting.     The   lungs   were    normal    except    for   a 

'  In  the  English  translation  of  Kiichenmeister's  work  on  Parasitology  (London,  1857). 
The  specimen  is  preserved  in  the  Hunterian  Museum,  London,  and  is  an  adult  liver  fluke, 
measuring  18  mm.  in  length  and  7  mm.  in  breadth. 


FASCIOLOPSIN^ 


245 


sharply  circumscribed  spot  at  the  base  of  the  left  lung.  Twenty 
days  later  during  a  fit  of  coughing  the  patient  spat  up  a  fluke  25  mm. 
long,  characterized  by  its  slender  aspect  and  by  the  size  of  its  ventral 
sucker,  and  its  close  proximity  to  the  oral  sucker.  Considering  the 
fact  that  Gouvea's  patient  had  spent  many  weeks  in  July  of  the  same 
year  in  Dakar  (Senegambia),  where  according  to  Railliet  Fasciola 
gigantica  is  common  in  slaughtered  animals,  and  considering  also 
the  characters  of  the  fluke,  Railliet  rightly  assumes  that  one  had  to 
do  with  the  African  giant  fluke  and  that  the  patient  had  infected 
himself  in  Dakar. 


Sub-family.     Fasciolopsinae,  Odhner,  1910. 
Genus.     Fasciolopsis,  Looss,   1898. 


Ventral  sucker  large,  and  elongated 
posteriorly  injto  a  sac.  Cirrus  pouch 
long  and  cylindrical,  its  greatest  length 
being  occupied  by  the  sinuous  tubular 
seminal  vesicle,  on  which  exists  a  peculiar 
caecal  appendage.   Laurer's  canal  present. 


Fasciolopsis  buski,  Lank.,  1857. 

Syn.  :  Distomuin  buski,  Lank.,  1857  ; 
Dist.  crassum,  Cobbold,  i860,  nee  v.  Sieb., 
1836. 

The  length  of  the  body  varies ; 
it  may  measure  24  to  37  or  even 
attain  70  mm.;  the  breadth  is  from 
5*5  to  12  to  14  mm.  In  the  pig  the 
fresh  parasites  measure,  smallest, 
12  to  8  mm. ;  largest,  35  to  16  mm. 
(Mathis  and  Leger).  Skin  without 
spines,  but  according  to  Heanly 
always  present  in  man  and  pig 
specimens.  The  oral  sucker 
measures  05  mm.  in  diameter; 
the  ventral  sucker  is  three  to  four 
times  as  large ;  the  pharynx  is 
globular,  0*7  mm.  in  diameter ; 
the  prepharynx  is  provided  with  a 
sphincter  ;  the  intestinal  caeca  ex- 
tend to  the  posterior  border  with 


Fig.  146. — Fasciolopsis  buski,  Lank.  F.j., 
ventral  sucker ;  C.p.,  cirrus  pouch  ;  /.,  inles- 
tinal  fork;  6'.2/.,  vitellaria ;  7'.,  testes;  0., 
ovary;  Ms.,  sucker;  Shg.,  shell  gland; 
6^/.,  uterus.     Magnified.     (After  Odhner.) 


246 


THE   ANIMAL  PARASITES   OF   MAN 


two  characteristic  curves,  one  at  the  anterior  border  of  the  anterior 
testis,  the  other  between  the  two  testes.  The  genital  pore  is  at  the 
anterior  border  of  the  ventral  sucker;  the  cylindrical  cirrus  pouch 
extends  from  behind  the  ventral  sucker  to  half-way  to  the  shell  gland. 
The  seminal  vesicle  extends  forwards  within  the  cirrus  pouch  as  a 
convoluted  tube.  From  its  anterior  portion  is  given  off  the  caecal 
appendage,  which  has  itself  short  lateral  diver- 
ticula. It  runs  backwards,  ending  blindly 
about  o*5  mm.  from  the  posterior  end  of  the 
cirrus  sac.  The  seminal  vesicle  is  continued  as 
the  pars  prostatica  (?)  0*5  mm.  long,  and  this 
by  the  very  short  ejaculatory  duct  (13  /x),  and 
finally  by  the  fairly  long  cirrus,  which  is  beset 
with  very  fine  spines  except  at  either  extremity. 
The  ovary  and  shell  gland  are  situated  at  about 
the  middle  of  the  body  with  the  testes  behind 
them,  and  the  uterus  in  front.  The  vitellaria 
extend  from  the  ventral  sucker  to  the  posterior 
border.  The  eggs  measure  120  fi  to  130 /i  in 
length  and  77  fiioSo  fi  in  breadth,  and  resemble 
those  of  Echinochasma  sp.  in  dogs.  The  larval 
stages  are  said  to  occur  in  shrimps. 
Habitat. — Intestine  of  pig  and  man. 
Distribution. — In  man  :  India,  Siam,  China, 
Assam,  Sumatra.  It  is  common  in  Cochin 
China  (16  out  of  133  Annamites,  Noc),  in 
Tonkin  very  rare.  Dr.  J.  Bell  has  sent  me 
[J.  W.  W.  S.]  human  specimens  from  Hong  Kong.  In  pigs  :  very 
common  in  South  China  (Heanly).  Common  in  pigs  in  Hong  Kong. 
Sixteen  out  of  248  pigs  (i.e.,  6  per  cent.)  infected  in  Hanoi. 


•  Fig.  147.  —  Fasciolopsis 
rathouisi^  Poir.  :  the 
mouth  at  the  top,  and  under 
it  the  genital  pore  and  ven- 
tral sucker,  behind  which 
again  is  the  uterus.  The 
vitellaria  are  at  the  sides, 
and  posteriorly  in  the  central 
field  the  ramified  testes  ;  the 
ovary  is  in  front  of  the  right 
testis.     (After  Claus.) 


Fasciolopsis  rathouisi,  Ward,   1903. 

Syn.  :  Distomum  rathouisi^  Poirier,  1887. 

Fifteen  to  19  mm.  long  by  8*5  to  10-5  mm.  broad  by  about  3  mm. 
thick.  Skin  with  spines  (Leiper).  Bluntly  oval  or  elliptical  with 
short  cephalic  cone  which  is  absent  in  Fasciolopsis  biiski.  Oral  sucker 
subterminal,  0*25  to  o'29  mm.  broad  by  0*2  mm.  in  antero-posterior 
diameter.  Distant  from  ventral  sucker  by  about  twice  its  diameter. 
Ventral  sucker  1-32  to  1*38  mm.  broad  by  0*68  to  07  mm.  in  antero- 
posterior diameter.  (Esophagus  extremely  short.  Cirrus  sac  not 
conspicuous  and  straight  as  in  Fasciolopsis  buski,  but  is  convoluted. 
Testes  one  behind  the  other  (according  to  Poirier  they  lie  beside  one 


FASCIOLOPSIS   FULLEBORNI  247 

another),  more  compactly  branched,  broader  and  denser  than  in 
Fasciolopsis  bnski.  Ovary  on  right  side,  small,  coarsely  branched. 
Uterus  in  broad,  closely  grouped  coils,  packed  with  ova  anterior  to 
ovary.  Vitellarian  acini  more  numerous  and  somewhat  differently 
distributed.  Eggs  150/^  by  80  yit,  thin  shelled.  [H.  B.  Ward,  who  has 
examined  this  species,  and  from  whose  account  the  above  is  mainly 
taken,  considers  that  it  is  a  good  species,  although  the  differences 
between  it  and  Fasciolopsis  buski  are  slight,  while  Odhner,  who  examined 
the  original  species,  is  of  the  opposite  opinion. — J.  W.  W.  S.]  The 
parasite  appears  to  cause  diarrhoea,  wasting  and  occasionally  jaundice. 

Habitat. — Intestine  of  man. 

Distribution. — China,  common  in  some  parts  (Goddard). 


Fasciolopsis  goddardi.  Ward,  1910. 

Twenty-one  to  22  mm.  long,  9  mm.  broad.  Skin  with  spines 
(Leiper).  Uterus  very  closely  coiled,  most  striking  character  is  the 
large  size  of  the  vitelline  acini.     Imperfectly  known. 

Distribution. — China  (Shanghai). 


Fasciolopsis  fulleborni,  Rodenwaldt,  1909. 

The  fully  extended  fluke  is  tongue-shaped,  50  by  14  mm.  ;  two 
contracted  specimens  measured  40  by  15  mm.  and  30  by  16  mm. 
respectively.  Skin  without  spines,  with  according  to  Leiper  cephaHc 
cone  not  clearly  defined.  Oral  sucker  circular,  075  mm.  in 
diameter,  slightly  larger  than  that  of  Fasciolopsis  buski.  Ventral  sucker 
2-6  mm.  in  diameter  (that  of  Fasciolopsis  buski  i*6  to  2  mm.).  Length 
2*9  mm.  (as  in  Fasciolopsis  rathoulsl),  the  excess  of  length  over  breadth 
being  due  to  the  posterior  elongated  sac-like  prolongation  of  the 
sucker.  Prepharyngeal  sphincter  present.  Pharynx  07  mm.  in 
diameter.  CEsophagus  practically  absent.  Gut  caeca  similar  to  those 
of  Fasciolopsis  buski. 

Testes — regularly  branched,  separated  by  an  incurving  of  the  caeca, 
the  anterior  occupying  a  smaller  area  than  the  posterior. 

Ovary — very  small,  as   in   Fasciolopsis    buski,    on    the   right   side. 

Shell  Gland — almond-shaped,  2-3  by  1*2  mm.  In  Fasciolopsis  buski 
it  is  round  and  smaller,  i  to  i"5  mm.  in  diameter. 

Vltellarla — similar  in  distribution  to  those  of  Fasciolopsis  buski,  but 
the  acini  are  strikingly  small. 

Cirrus  Sac — is  the  most  characteristic  feature  of  this  species.  It  is 
a  powerfully  built,  convoluted  sac  standing  out  clearly  on  the  body. 


248 


THE   ANIMAL   PARASITES   OF   MAN 


t^* Oral  sucker 

■  Prepharynx 

Pharyny 


':^*^    Vl'A7fra/  sucker 


Vagina 


Cut 

Cirrus  sac  (with 
seminal  vesicle) 

'  Uterine  coils 


-  Receptaculum  seminis 

"  Shellgland  and  yolk 
receptacle 

"~  Ovary 
---Gut 

r:-  Testes 


l--;-'  Excretory  system 


y  Vitellaria 


Excretory  pore  (Dorsal) 

Fig.  i/\%.—Fasciolopsi5fulleborni,  ventral  aspect.     {After  Fiilleborn.) 


TROGLOTREMID^  249 

It  is  not  a  uniform,  straight  cylinder  0*25  to  0*33  mm.  in  diameter,  as 
in  Fasciolopsis  hiiski,  but  even  in  fully  extended  flukes  is  typically 
convoluted.  It  is  i  mm.  thick  in  the  middle,  but  in  other  parts  varies 
much  from  this.  The  posterior  end  of  the  cirrus  sac  is-  at  two-thirds 
or  more  of  the  distance  from  ventral  sucker  to  shell  gland.  In  the 
case  of  Fasciolopsis  bnski  the  posterior  end  of  the  sac  only  extends 
half-way. 

Seminal  Vesicle — has  a  peculiar  convoluted,  saccular  and  angular 
course,  but  the  caecal  appendage  characteristic  of  the  genus  appears 
to  be  absent  ! 

Excretory  System. — The  main  stem  gives  off  very  regular  transverse 
branches  which  are  well  seen  posteriorly. 

Eg^s. — 100  //,  by  73  //,.     Thin  shelled. 

Habitat. — Intestine.     Mahommedan  from  Calcutta. 

[It  is  evident  that  a  re-examination  of  fresh  material  is  required 
before  the  validity  of  all  these  species  can  be  accepted. — J.  W.  W.  S.] 


Family.     Troglotremidae,  Odhner,  1914.  ' 

Genus.     Paragonimus,  Braun,  1899. 

Body  egg-shaped  or  somewhat  elongated,  generally  more  broadly  rounded  in 
front  than  behind.  Covered  all  over  with  spear-shaped  spines  arrangedin  groups. 
Gut  caeca  winding  with  dilatations  or  constrictions  in  parts.  Ventral  sucker  in  or 
in  front  of  the  middle  of  the  body.  Excretory  bladder  cylindrical,  very  long  and 
broad,  reaching  in  front  to  the  bifurcation  of  the  gut.  The  lateral  excretory  canals 
join  the  bladder  only  a  little  in  front  of  the  excretory  pore.  Genital  pore  median 
just  behind  the  ventral  sucker.  Genital  sinus  duct-like.  Cirrus  sac  absent.  Male 
terminal  organs  very  small.  Ejaculatory  duct  present.  Testes  and  ovary  deeply 
lobed,  the  testes  in  or  just  behind  the  middle,  the  ovary  somewhat  laterally  placed 
just  behind  the  ventral  sucker.  Uterus  forms  a  coil  behind  the  ventral  sucker. 
Eggs  rather  large,  thin  shelled,  the  ovarian  cell  still  unsegmented  on  deposition. 
Receptaculum  seminis,  small. 

Parasitic  in  the  lungs  of  mammals,  enclosed  in  cyst-like  cavities,  generally  in 
pairs. 

Type  species. — P.  westermajiii  in  the  tiger. 


Paragonimus  ringeri,'  Cobb.,  1880. 

Syn.  :  Distoma  ringeri,  Cobb.,  1880;  Distoma  pulmonale.,  Baelz,  1883  ; 
Distoma  pulmo7iis,  Suga,  1883. 

The  body  is  of  a  faint  reddish-brown  colour  and  plump  oval 
shape.  The  ventral  surface  a  little  flattened;  7*5  to  12  mm.  in  length, 
4  to  6  mm.  in  breadth,  and  3-5  to  5  mm.  thick  (in  man).  The  oral 
sucker  (075  mm.)  is  subterminal;  the  ventral  sucker  (o*8  mm.) 
somewhat  in   front  of  the  middle    of  the  body.     Pharynx  spherical, 


250 


THE   ANIMAL   PARASITES   OF   MAN 


0 


0*3  mm.  in   diameter,  or  0*4  by  0*3    mm.  ;    oesophagus,  0*02    mm.  ; 

intestinal  caeca  convoluted,  asymmetrical,  the   first   part   having   the 

same  structure  as  the  oesophagus.  The 
cuticle  is  covered  with  spines  in  groups  ; 
the  excretory  pore  opens  at  the  posterior 
end  rather  on  the  ventral  surface,  the  excre- 
tory ducts  open  into  the  elongated  bladder 
at  the  hind  end  near  the  pore.  Genital 
pore  behind  •  the  ventral  sucker  and 
median.  Genital  sinus  0*2  mm.  long 
with  thick  wall,  ejaculatory  duct  o'i3  mm., 

pars  prostatica  0*2  mm.,  seminal  vesicle  duct-like  of  irregular  outline. 

Behind    the    sucker   the  ovary  on    the   left,  and  the  closely  packed 

uterine  coil  on  the  right  (though  amphitypy  of  these  two  organs  is 

common) ;  the  two  irregularly  lobed  testes  lie  side  by  side  posteriorly. 

Vitellaria  extensive,  leaving  only  a  median  dorsal  and  ventral  space 


Fig.  149. — Paragonimus 
ringeriy  Cobb. :  to  the  right, 
dorsal  aspect ;  to  the  left,  ventral 
aspect.  Natural  size.  (After 
Katsurada.) 


Fk;,  150. — Paragonimus  ringeri,  Cobb.  : 
diagram  of  the  internal  organs.  a,  oeso- 
phagus ;  3,  vitellaria  (a  portion  only  shown) ; 
f,  common  genital  duct ;  d,  shell  gland  with 
oviduct,  Laurer's  canal  and  vitelline  duct ;  e, 
ovary;  y,  vitelline  receptacle;  g,  excretory 
pore  ;  A,  oral  sucker ;  i,  pharynx  ;  k^  gut*;  /, 
ventral  sucker;  w,  uterine  coils;  «,  vitellarian 
ducts ;  (7,  vas  efferens ;  /,  testis.  (After  Kubo. ) 


Fig.  150A. — Paragonimus  wesiermanii. 
Kerb.  :  seen  from  the  ventral  surface. 
Mouth,  pharynx,  intestinal  caeca,  at  the  sides 
of  which  the  vitellaria  are  observed.  The 
genital  pore  is  behind  the  ventral  sucker,  and 
next  to  it,  on  the  left,  the  ovaty  ;  on  the  right, 
the  uterus  ;  the  two  testes  posteriorly  ;  the 
excretory  vessel  in  the  middle.  lo/l.  (After 
Leuckart.) 


free.  Seminal  receptacle  probably  absent ;  Laurer's  canal  present. 
The  eggs  are  oval,  brownish-yellow,  fairly  thin  shelled,  and  measure 
onf-an  average  81 '2  /a  by  49*2  /x. 

The   following  species   are  also   known  : — P.  westennaiiii,  Kerb., 
1878,  in  the  tiger,  and  P.  kellicoiti,  Ward,  1908,  in  the  pig,  dog,  and 


PARAGONIMUS   RINGERI 


251 


cat  (N.  America).     Ward  and  Hirsch  give  the  following  differences 
between  the  spines  of  the  three  forms  : — 


P.  ringeri. 

P.  westermanii. 

P.  kellicotti. 

Shape 
Distribution 

Chisel-shaped,  mod- 
erately heavy. 
Circular  rows,  in 

Lancet-shaped, 

very  slender. 

Circular  rows. 

Chisel-shaped, 

heavy. 
Circular  rows, 

groups. 

in  groups. 

singly. 

Two  other  species,  P.  riidis,  Diesing,  1850,  in  a  Brazilian  otter 
{Ltitra  brasiliensis)f  and  P.  compachis,  Cobbold,  1859,  in  the  Indian 
ichneumon,  are  but  little  known. 

Habitat. — Lungs,  pleurae,  and  especially  the  bronchi  of  man  and 
dog.  The  alleged  occurrence  (of  eggs)  in  other  organs  may  be  due 
to  confusion  with  those  of  Schistosoma  japoniciim. 

Distribiition. — China,  Korea,  and  especially  in  Japan,  where,  accord- 
ing to  Katsurada,  there  are  no  districts  that  are  entirely  free  from  pul- 
monary flukes.     The  mountainous  provinces  of  Okayama,  Kumamoto, 
Nagano    and    Tokushima   are    the    principal 
centres. 

Pathology.  —  The  number  present  in  the 
lung  varies  from  two  to  twenty,  about. 
Usually  one  cyst  contains  one  worm,  but  in 
the  dog  each  cyst  contains  two.  The  cysts 
admit  the  tip  of  the  finger,  and  have  a  fibrous 
wall  I  mm.  thick.  They  origin^ate  partly  from 
dilatation  of  bronchi  and  bronchioles.  Others 
arise  from  the  inflammatory  reaction  of  lung 
tissue  into  which  the  worms  have  wandered. 
The  worms  and  their  eggs  cause  bronchitis 
and  peribronchitis,  catarrhal,  haemorrhagic,  or 
purulent,  and  areas  of  consolidation.  Areas 
containing  eggs  in  their  centre  resembling 
tubercle  nodules  are  not  uncommon,  and 
extensive  cirrhosis  of  the  lung  may  be  found. 

As   a  result  of   these  changes,    emphysema   and   bronchiectasis  also 
occur. 

As  to  the  development,  only  the  following  details  are  known  : 
that  the  eggs,  which  before  segmentation  of  the  ovum  reach  the  open 
in  the  sputum  and  through  being  swallowed  also  in  the  faeces, 
develop  in  water  into  a  miracidium  ciliated  all  over,  which  hatches 
and  swims  about  freely.  According  to  Manson  this  takes  place  in 
four  to  six  weeks. 


Fig.  151. — Egg  of  Para- 
gonivius  ringeri,  Cobb. , 
from  the  sputum.  Showing 
the  ovarian  cell  and  vitel- 
line cells  and  granules. 
1,000/1.    (After  Katsurada.) 


252  THE   ANIMAL   PARASITES   OF   MAN 

Sub-family.     Opisthorchiinae,  Looss,  1899. 

Genus.     Opisthorchis,  R.  Bl?.nch.,  1845. 

OpisthorchiinjE  with  lobed  testes.  Laurer's  canal  present.  Parasitic  in  the  bile- 
ducts  of  mammals  and  birds. 

Opisthorchis  felineus,  Riv.,  1885. 

Syn. :  Distoma  co7ius,  Gurlt,  1831  {tiec  Creplin,  1825);  Distoma  lanceolatum^ 
V.  Sieb.,  1836,  V.  Tright,  1889  {?iec  Mehlis,  1825  =  Fasciolo  lanceolata^Kud.^  1803); 
Distoma  sibiricum.  Winogr.,  1892  ;  Distoma  tenuicolle,  Miihl.,  1896. 

This  parasite  is  yellowish-red  in  the  fresh  condition,  and  almost 
transparent.  The  body  is  fiat,  with  a  conical  neck  at  the  level  of  the 
ventral  sucker  marked  by  a  shallow  constriction  ;  this,  however, 
is  only  noticeable  in  fresh  and  somewhat  contracted  specimens. 
Posteriorly  to  the  ventral  sucker  the  lateral  borders  run  fairly  parallel ; 
the  posterior  end  is  either  pointed  or  rounded  off.  The  length  and 
breadth  vary  according  to  the  contraction,  being  usually  8  to  11  mm. 
by  1*5  to  2  mm.  The  suckers  are  about  one-fifth  to  one-sixth  of 
the  length  of  the  body  distant  from  each  other,  and  of  about 
equal  size  (o'23  to  0*25  mm.).  The  oesophagus  is  hardly  any  longer 
than  the  pharynx,  which  lies  close  behind  the  oral  sucker ;  the 
intestinal  caeca  reach  almost  to  the  posterior  border  and  are  often 
tilled  with  blood.  The  excretory  pore  is  at  the  posterior  extremity, 
and  the  excretory  bladder  forks  in  front  of  the  anterior  testis.  The 
testes  in  the  posterior  fourth  of  the  body  lie  obliquely  one  behind 
the  other  ;  the  anterior  one  has  four  lobes,  the  posterior  one  five 
lobes ;  the  ovary  is  in  the  median  line  transversely,  simple  or 
slightly  lobed ;  behind  it  lies  the  large  pear-  or  retort-shaped  recepta- 
culum  seminis  and  Laurer's  canal.  The  uterus  is  in  the  median  field. 
The  vitellaria  occupy  the  fairly  broad  lateral  areas,  in  about  the 
central  third  of  the  body,  beginning  behind  the  ventral  sucker  and 
terminating  at  about  the  level  of  the  ovary ;  the  acini  are  small  and 
arranged  in  groups  of  seven  to  eight,  separated  by  interstices.  The 
genital  pore  is  close  in  front  of  the  ventral  sucker.  The  ^ggi^  are 
oval  with  sharply  defined  operculum  at  the  pointed  pole,  30  yit  by  1 1  yit. 

This  species,  which  is  frequently  confused  with  others,  inhabits  the  gall- 
bladder and  bile-ducts  of  the  domestic  cat  especially  ;  but  is  also  found  in  the 
dog,  in  the  fox,  and  in  the  glutton  {Gulo  borealis).  It  has  been  observed  in 
France,  Holland,  North  Germany  (being  particularly  frequent  in  East  Prussia),  in 
Russia,  Scandinavia,  Siberia,  Japan,  Tonkin,  Hungary,  and  Italy.  The  North 
American  form  (from  cats  and  Ca7iis  latrans)  is  a  distinct  species  {Opisthorchis 
pseudofelineus) . 

In  man  this  species  was  first  found  by  Winogradoff  in  Tomsk  (nine 
cases),  then  by  Kholodkowsky  in  a  peasant  from  the  neighbourhood 


OPISTHORCHIS    FELINEUS 


253 


of  Petrograd  who  had  travelled  a 
great  deal  in  Siberia,  and  finally  by 
Askanazy  in  five  persons  who  were 
natives  of  the  East  Prussian  district  of 
Heydekrug.  In  Tomsk,  Opisthorchis 
fellnetis  is  the  most  frequent  parasite 
of  man  that  comes  under  observation 
^t  post  inorteni  (6*45  per  cent),  whereas 
Tcviiia  sagiuafa  has  only  been  found 
in  3*2  per  cent.,  Echinococcif^  in  2*4  per 
cent.,  Ascaris  himbricoides  in  i'6  per 
cent.,  and  Oxyuris  vennicidaris  in  0*8 
per  cent,  of  the  autopsies.  In  the  district 
of  Heydekrug,  however,  the  species  in 
question  is  also  frequent,  as  in  a  few 
years  five  cases  came  to  our  knowledge 
(of  which  three  were  diagnosed  by  the 
discovery  of  the  eggs  in  the  faeces). 

In  none  of  Winogradoff's  nine  cases 
had  the  death  of  the  patient  been  caused 
direct  by  the  parasites,  yet  more  or  less 
extensive    changes    in    the    liver    were 
found  in  all  of  them;  such  as  dilatation 
of  the  bile-ducts  with  inflammation  and 
thickening  of  their  walls,  and  foci  of 
inflammation    or   atrophy  in  the  liver 
substance  ;    icterus    was    present    five 
times  and  atrophy  of  the  liver  an  equal 
number  of  times ;  ascites 
was  observed  three  times, 
and  in  two  cases,  probably 
of  recent  date,  the  organ 
was  enlarged.     The  num- 
ber    of     parasites     found 
fluctuated  between  a   few 
and  several  hundreds. 

In  two  of  Askanazy's 
cases,  which  he  examined 
more  closely,  carcinoma 
which  had  developed  at 
the  places  most  invaded  by  flukes  was  found  at  the  posUmortem.,  so 
that  perhaps  there  may  be  grounds  for  the  connection  which  the  author 
seeks  to  establish  between  cancer  of  the  liver  and  the  changes  induced 
by  the  parasites  ;  these  changes  consist  of  numerous  and  even  ramified 


'^^ 


r.s. 


Fig.  152.— 
Egg  of  Opis- 
thorchis feli- 
neus,  Riv. 
830/1. 


Fig.  153.  —  Opisthorchis  felineus  : 
from  the  cat.  m.,  mouth;  p.6., 
pharynx  ;  i.,  gut ;  ^./.,  genital  pore  ; 
ac,  ventral  sucker  ;  ut.,  uterus  ;  v.g., 
vitellarium ;  ov.,  ovary;  s.g.,  shell 
gland;  r.s.,  receptaculum  seminis ; 
t.  testes ;  ex.  /.,  excretory  pore. 
(After  Stiles  and  Hassall.) 


254 


THE  ANIMAL   PARASITES   OF   MAN 


m 


s.g. 


L.c. 


t. 


7lt. 


v.g. 


v.df. 


r.S. 


/ 


ex.  c. 


ex.  p. 


proliferations  of  the  epithelium  of  the 
biliary  duct  into  the  connective 
tissue,  which  is  likewise  proliferated. 
The  number  of  worms  found  in  one 
case  amounted  to  over  lOO ;  in  a 
second  case,  in  which  the  parasites 
had  also  invaded  the  pancreatic  duct, 
their  number  was  even  larger. 

Winogradoff  as  well  as  Askanazy 
found  isolated  flukes  in  the  intestine 
also. 

Unfortunately,  nothing  much  is 
known  of  the  history  of  the  develop- 
ment of  Opisthorchis  felmeus ;  we  only 
know  that  when  deposited  the  eggs 
already  contain  a  ciliated  mira- 
cidium,  which,  however,  according 
to  my  experience,  does  not  hatch  out 
in  water,  but  only  after  the  entry  of 
the  eggs  into  the  intestine  of  young 
Limriceus  stagnalis  ;  no  further  de- 
velopment, however,  occurs.  Wino- 
gradoff states  that  he  has  seen  the 
miracidia.  hatch  after  the  eggs  had 
been  kept  in  water  for  a  month  at 
37°  C. ;  and  has  even  observed  free 
miracidia  in  the  bile  of  man  and  of 
a  dog  respectively.  Although  the 
whole  post-embryonal  development 
of  the  cat  fluke  remains  yet  to  be 
investigated,  Askanazy  by  a  series  of 
experiments  on  cats  and  dogs  has  dis- 
covered the  mode  of  infection.  The 
intermediate  hosts  are  fish,  and 
mainly  the  ide,  in  this  country  called 
Tapar  (Idus  melanoiuSj  H.  and  Kr.), 
and    of    subsidiary    importance    the 


Fig.  154. — Opisthorchis  psiudofdineus  :  from  the 
bile-duct  of  the  cat  (Iowa),  w.,  oral  sucker;  p.h.^ 
pharyngeal  bulb  ;  ^j.,  oesophagus  ;  i.,  intestine;  va.^ 
vagina;  g.p.m.,  male  orifice;  ac,  ventral  sucker; 
«/.,  uterus  ;  v.g.,  vitellarium  ;  s.g.,  shell  gland  ;  v.dt.^ 
vitelline  duct ;  ov. ,  ovary ;  r.s. ,  receptaculum  seminis  ; 
L.c,  Laurer's  canal;  /.,  testis;  ex.c,  excretory 
bladder;  ex.p.,  excretory  pore.     (After  Stiles.) 


PAROPISTHORCHIS 


255 


roach  (Leiiciscus  rutiltis).  Both  species  of  fish  as  well  as  others  are 
readily  eaten  raw  by  man  on  the  Courland  lagoon  (Baltic).  It  is,  more- 
over, significant  that  those  persons  whom  Askanazy  found  infected 
with  the  cat  fluke  were  also  infected  with  Dihothriocephaliis  latus,  the 
intermediate  host  of  which  is  also  fish  (Lota  sp.,  Esox  sp.,  Perca  sp.). 

In  one  of  his  nine  cases  Winogradoff  also  saw  a  small  fluke 
covered  all  over  with  spines,  which  he  conjectured  to  be  the  young 
stage  of  Opisthorchis  felineiis ;  as,  however,  according  to  my  experi- 
ence, this  species,  even  in  smaller  specimens,  is  always  without  spines, 
the  above  hypothesis  cannot  be  accepted.  It  is  much  more  probable 
that  one  of  the  other  species  that  also  invade  the  liver  of  cats  may 
accidentally  be  introduced  into  man;  we  know,  in  fact,  that  Metorchis 
albidtis,  Braun,  and  Metorchis  truncatus,  Rud.,  are  both  covered  with 
spines.  As,  however,  the  spines  of  the  first-named  species  are  rather 
apt  to  fall  off,  and  also  as  it  possesses  a  different  shape  (spatula-shaped), 
it  may  be  assumed  that  probably  Winogradoff  had  found  Metorchis 
truncatiis,  Rud.,  18 19,  in  his  patient. 

Genus.     Paroplsthorchis,  Stephens,  1912. 

Structure  as  in  Opisthorchis,  except  that  the  ventral  sucker  and  genital  pore  occur 
on  the  apex  of  a  process  or  pedicle  projecting  from  the  anterior  portion  of  the  body. 
This  process  is  about  ^  mm.  long,  and  is  retractile. 

Paroplsthorchis   caninus,  Barker,  1912. 

Syn.  :  Distoma  co?ijunctum^  Lewis  and  Cunningham,  1872  ;    Opisthorchis  noverca^ 
M.  Braun,  1903  (^pro  parte)  ;  Opisthorchis  caninus^  Barker,  19 12  (?). 

Length  varies  from  275  to  575  mm.  in  preserved  specimens,  average 
3*6  to  5'2  mm.  Body  uniformly  spinose,  though  as  a  rule  spines  are 
not  present  on  the  pedicle.  Body  slightly  concavo-convex,  the  con- 
cavity being  ventral.  Oral  sucker  0*28  mm.  Pharynx  0*224  by 
0'i84  mm.  (Esophagus  0*04  mm.  Ventral  sucker  0"i76  mm.  in 
diameter.     Pedicle  about  \  mm?  long,  may  be  completely  retracted. 

Genital  Pore — opens  on  the  apex  of  the  pedicle  in  front  of  the 
ventral  sucker.  Its  exact  position  varies  with  the  state  of  contraction 
of  the  parts.  In  certain  cases  it  actually  opens  within  the  cuticular 
border  of  the  sucker,  in  other  cases  it  opens  externally  to  the  sucker 
and  anterior  to  it.  The  opening  is  covered  with  scales.  The  vas 
deferens  and  uterus  run  alongside  one  another  until  they  merge  near 
the  apex  of  the  pedicle  into  a  common  sinus. 

Vitellaria — consist  of  eight  acini  on  each  side,  extending  from 
slightly  behind  the  base  of  the  pedicle  to  the  anterior  border  of  the 
ovary,  or  as  far  back  as  a  line  separating  the  posterior  border  of  the 
ovary  from  the  anterior  border  of  the  anterior  testis. 


256 


THE   ANIMAL   PARASITES   OF   MAN 


V.  .ex.  lat 
V.  seui. 


P.  ex. 


Fig.  155. — Paropisthorchis  caninus :  from  the  bile-ducts  of  the  pariah  dog,  India.  AceL  v.y 
ventral  sucker  ;  £//.,  uterus  ;  V.  ex.  /a/.,  longitudinal  excretory  duct ;  V.  sem.,  seminal  vesicle; 
Se?n.  r«<r.,  seminal  receptacle;  Ov.,  ovary;  V.  «:*:.,  excretory  bladder;  TesL  /.,  left  testis; 
7<!j/.  r.,  right  testis ;  /*.  ^jr.,  excretory  pore,      x  40.    (After  Stephens.) 


AMPHIMERUS 


257 


-ac- 


v.g 


Testes. — Anterior  testis  0*496  by  0*44  mm. ;  posterior  testis  0*52  by 
0*48  mm.,  usually  ovoid,  though  both  may  be  regularly  lobed.  The 
anterior  testis  is  usually  on  the  left  side. 

Ovary — multilobular,  the  lobes  6  to  8  being  irregular  in  size  and 
shape. 

Shell  Gland — extensive  and  diffuse,  occupying  an  area  which  approx- 
imately corresponds  with  the  loop  of  the  transverse  vitelline  ducts. 

Seminal  Receptacle — globular,  to  the  right  of  and  dorsal  to  the 
posterior  lobe  of  the  ovary. 

Laurel's  Canal  —  generally 
runs  from  the  end  of  the  re- 
ceptacle with  a  single  curve 
medially  and  backwards. 

Uterine  Coils — form  loosely 
packed  transverse  coils  ter- 
minating slightly  in  front  of 
the  level  of  the  first  vitelline 
acini.  From  here  the  uterus 
passes  forwards  into  the  pedicle 
to  the  left  and  ventral  to  the 
seminal  vesicle. 

Seminal  Vesicle  —  com- 
mences about  the  level  of  the 
first  vitelline  acini.  The  coils 
displace  the  uterus  ventrally  and 
to  the  left.  In  the  pedicle  the 
vesicle  diminishes  in  extent  and 
lies  in  its  dorsal  (anterior)  side. 

Habitat. — Liver  of  pariah 
dogs,  India.  In  North-Western 
Provinces  about  40  per  cent, 
are  infected.  This  fluke  ap- 
pears to  be  different  from 
Amphimerus  {Op isth orchis) 
noverca  in  man,  as  the  latter  has 
not  the  pedicle  on  the  summit 
of  which  lie  the  sucker  and 
common  genital  pore. 


v.g. 


Genus.  Amphimerus,  Barker, 

1912  (?). 
Structure  as  in  Opisthorchis,  ex- 
cept that  the  vitellaria  are  separated 
into  two  portions,  an  ant-ovarial  and 
a  post-ovarial. 
17 


X)V. 


-y.d. 


«x.o. 


Fig.  \^^.^ Amphimerus  noverca,  BiSLun.i  ,::o.(94 
oral  sucker  ;  p.b.,  pharynx;  ac,  ventral  sucker  ; 
ut.,  uterus;  v.g,,  vitellarium  ;  ov.,  ovary;  v.d.,  vas 
efferens;  ex. c,  excretory  canal ;  t.,  testis.  (After 
McConnell.) 


258 


THE   ANIMAL   PARASITES   OF   MAN 


Ms. 


Amphimerus  noverca,  Barker,  191 2  (?)• 
Syn.:  Distommn  conjiinctum,  McConnell,  1876  {nee  Cobbold,  1859);  Opisthorchis 
noverca,  M.  Braun,  1903  pro  parte. 
At  the  autopsy  of  two  Mahommedans  who  died  in  Calcutta, 
McConnell  found  a  large  number  of  Distomata  in  the  thickened  and 
dilated  bile-ducts.  The  worms  were  lancet-shaped,  covered  with 
spines,  and  measured  9-5  to  127  mm.  in  length  and  2*5  mm.  in  breadth. 
The  two  suckers  lie  very  close  to  one  another,  the  anterior  one  bemg 
larger  than  the  ventral ;  the  genital  pore  opens  immediately  in  front 
of  the  ventral  sucker  ;  pharynx  spherical ;  intestinal  caeca  extending  far 
back.  At  the  commencement  of  the  posterior  third  of  the  body  the 
two  testes,  somewhat  apart,  the  anterior  one  roundish,  the  posterior  one 

distinctly  lobed.  The  transverse 
and  slightly  lobed  ovary  in  front 
of  the  bifurcation  of  the  Y-shaped 
excretory  bladder,  whence  the 
uterus,  in  convolutions  barely 
spreading  beyond  the  central 
field,  extends  to  the  pore;  the 
vitellaria  in  the  lateral  areas 
commence  behind  the  ventral 
sucker  and  extend  to  the  testes. 
Cirrus  pouch  absent.  Eggs  oval, 
34  /x  by  21  /^. 

Genus.  Clonorchis,  Looss,  1907. 

Structure  as  in  Opisthorchis,  dis- 
tinguished, however,  by  the  branched 
testes  situated  one  behind  the  other, 
the  branches  of  which  ventrally  en- 
croach upon  the  gut  forks  ;  dorsal 
to  the  testes  the  S-shaped  excretory 
bladder,  the  main  branches  of  which, 
arising  at  the  level  of  the  bifurcation 
of  the  gut,  open  into  the  bladder  below 
its  anterior  end.  Parasitic  in  the  bile- 
ducts  of  mammals  and  man. 


Vsc. 


Ex. 


Fig.  157.  —  Metorchis  conjmicius,^  (Syn.: 
Distomum  conjunctum,  Cobb.,  nee  Lew.  and 
Cunn.,  nee  McConn.)  :  from  Cants  fulvus. 
Fj.,  ventral  sucker;  /.,  intestine;  Vsc.y  vitel- 
laria ;  Ex.^  excretory  bladder ;  7".,  testes  ; 
(?.,  ovary;  Ms.,  oral  sucker;  Ph.^  pharynx; 
Ut.,  uterus.     (After   Cobbold.) 


Clonorchis   sinensis,  Cobbold,  1875. 

Syn.:    Distoma  sinense,  Cobbold,  1875;   Distoma  spathulatum.,  R.  Leuckart,  1876 
{nee  Rudolphi,  1819) ;  Distoma  hepatis  innoeuuin,  Baelz,  1883. 

In  shape  resembles  Opisthorchis  felineus,  13  to  19  mm.  long,  3  to 
4  mm.  broad,  at  the  beginning  of  sexual  maturity  12  to  13  mm.  long,  2*5 
to  3  mm.  broad.     Oral  sucker  o'58  to  0*62  mm.,  ventral  sucker  0*45  to 

'  This  species  from  Canis  fulvus  was  for  long  thought  to  be  the  same  as  that  here  described 
as  Amphimerus  noverca.    It  probably  does  not  belong  to  the  genus  Metorchis. 


CLONORCHIS   ENDEMICUS 


259 


0-49  mm.  in  transverse  diameter.  In  the  parenchyma  numerous 
yellowish  or  brownish  granules,  especially  behind  the  oral  sucker  and 
at  the  posterior  end.  Testicular  branches  very  long,  in  the  anterior 
testis  often  four,  in  the  posterior  testis  five  branches.  Ovary  generally 
with  three  large  lobes  and  a  smaller  lobe.  Vitellaria  not  always  sym- 
metrical, generally  extending  laterally  from  the  ventral  sucker  to  the 
ovary,  interrupted  in  parts. 

Eggs  26  fi  to  30  //,  by  15  fjL  to  17  fi. 
Average  29  /x  by  16  /^. 

This  (?)  species  was  discovered  in  1874 
by  McConnell,  in  Calcutta,  in  the  bile-ducts 
of  a  Chinaman  who  died  shortly  after  being 
admitted  into  hospital. 

Habitat. — Bile-ducts  of  man,  dog  and  cat. 

Distribution. — Especially  in  China,  ap- 
parently rare  in  Japan. 

Clonorchls  endemicus,   Baelz,  1883. 

Syn.  :  Disioma  sinense  s.  spathulatum  p.p. ; 
Distoma  hepatis  endemicum  s.  perniciosum,  Baelz, 
1883;   Distoma  japonicu7n,  R.  Blanchard,  1886. 

Very  similar  to  the  previous  species  and 
consequently  generally  confused  with  it. 
Length  between  6  and  13  mm.,  width 
varying  between  i'8  and  2*6  mm.  Oral 
sucker  0*37  to  0*5  mm.,  usually  0*43  to 
o*45  mm.  in  transverse  diameter ;  ventral 
sucker  0*33  to  0*45  mm.,  usually  0*37  to 
0*40  mm.  No  pigment  in  parenchyma  ; 
anterior  testis  with  four,  posterior  testis  with 
five  branches.     Vitellaria   continuous,  ova 

26  /X  by   13  yLt  to   16  yLfc. 

Habitat. — Bile-ducts  of  man,  dog,  cat 
and  pig. 


Ex 


\^ 


Fig.  158. — Clonorchis  sinensis. 
C.L.y  Laurer's  canal ;  Dst.,  vitel- 
laria; Ex.,  excretory  bladder; 
H.,  testes  ;  A'.,  ovary  ;  K.s.,  re- 
ceptaculum  seminis  ;  Vd.,  ter- 
minal section  of  vas  deferens. 
Magnified  4^  times.  (After 
Looss.) 


P'iG.  159. — Ova  of  Clonorchis  sinensis.     The  knobs  on  the  ends  of 
the  eggs  are  not  shown.     900/1.     (After  Looss.) 


26o 


THE   ANIMAL   PARASITES   OF   MAN 


Vitellarium 


Ovary 


Testis 


Distribution.— This  species  occurs  very  frequently  in  man,  in 
certain  districts  of  Japan,  especially  in  the  province  of  Okayama, 
Central  Japan,  in  particular  localities  of  which  above  60  per  cent,  of 
the  population  are  infected.  The  worms  are  sometimes  found  in 
enormous    numbers    in    the   liver    (upwards    of   4,000),    also   in   the 

pancreas  and  rarely  in  the  duo- 
denum. It  is  common  in  Tonkin 
and  Indo-China.  Leger  in  Tonkin 
found  50  per  cent,  of  people  ap- 
parently in  normal  health  infected, 
so  that  probably  symptoms  only 
arise  when  the  infection  is  intense. 
[The  exact  distribution  of  these  two 
species  is,  however,  not  precisely 
defined  at  present,  as  commonly  no 
distinction  is  made  between  them. — 
J.  W.  W.  S.] 

Verdun  and  Bruyant  deny,  in 
opposition  to  Looss,  the  possibility 
of  being  able  to  distinguish  within 
the  genus  Clonorchis  the  two  species 
described,  but  they  admit  the  justi- 
fication for  the  new  genus.  They 
also  report  the  occurrence  of  Opis- 
thorchis  felineus  in  man  in  Tonkin 
(Compt.  Rend.  Soc.  de  Biol.y  Ixii,  1907). 
Pathology. — Both  species  of  Clon- 
orchis give  rise  to  grave  symptoms.  The  liver  is  generally  enlarged, 
though  when  the  infection  has  lasted  some  time  it  begins  to  contract. 
The  surface  of  the  organ  is  studded  with  white  vesicles,  and  on  cutting 
into  it  one  sees  numer- 
ous cavities  with  thick- 
ened walls  (distended 
bile-ducts)  filled  with  a 
brownish  fluid  contain- 
ing innumerable  eggs, 
which  cause  its  colour. 
Microscopically,  the  epi- 
thelium of  the  bile-ducts 
is  either  (i)  entirely  de- 
stroyed, or  (2)  actively 
proliferates,  forming  an  adenomatous  outgrowth.  Occasionally  this 
proliferation  is  not  limited  by  the  wall  of  the  bile-duct  but  pene- 
trates it  and  leads  to  a  growth  of  numerous   new  ducts,  forming  a 


Seminal 
receptacle 

Excretory 
vesicle 


Fig.  160. — Clonorchis  endemicus.     x  6 
about.     (After  Looss.) 


Fig.  161. — Clonorchis  endemicus  :  eggs.     The  knobs 
on  the  eggs  are  not  shown,     x  900.     (After  Looss.) 


METORCHIIN^  261 

malignant  biliary  adenoma.  The  bile-ducts  have  their  connective 
tissue  wall  greatly  sclerosed.  These  fuse  with  one  another,  forming 
areas  of  sclerosis  devoid  of  liver  tissue.  As  a  result  of  these  changes 
the  liver  cells  atrophy  and  undergo  fatty  pigmentary  and  granular 
degeneration.  Besides  these  changes,  due  probably  to  the  toxic  action 
of  the  flukes,  mechanical  obstruction  due  to  the  actual  plugging  of  the 
ducts  by  the  flukes  causes  retention  of  bile  and  icterus,  and  through 
pressure  on  veins,  ascites  and  hypertrophy  of  the  spleen. 

To  what  extent  blood  or  bile  respectively  forms  the  food  of  the 
flukes  is  uncertain. 

Life-history. — (Kobayashi,  191 1,  Mitteihingen  aiis  dein  kaiscrlichen 
InstiUit  jilr  Infektions-Krankheiten  zii  Tokio,  pp.  58-62.) 

It  results  from  the  work  of  Kobayashi  in  Japan  that  fresh-water  fish 
form  the  second  intermediate  host  for  Clonorchis  endemicus.  He  fed 
cats  with  encysted  flukes  (cercarise)  from  various  fish  and  easily 
succeeded  in  infecting  them,  e.g.  a  kitten,  proved  to  be  uninfected  by 
repeated  examination  of  its  faeces,  was  fed  on  infected  fish  ;  a  month 
later  innumerable  flukes  were  found  in  the  bile-ducts,  gall-bladder, 
pancreas  and  even  in  the  duodenum.  The  fish  infected  were  Leuco- 
gobis  giintheri,  Pseudorashora  parva,  and  to  a  less  extent  A checlognathtis 
lanceolata,  Acheclognathus  limbata,  Paracheclognathus  rJionibea,  Pseudo- 
perilampns  typiis,  Abboltina  psegma,  Biwia  zezera  and  Sarcocheilichihys 
variegatus.  The  cysts  occur  throughout  the  muscles  and  subcutaneous 
tissue  of  the  fish.  Length  0*13  mm.,  breadth  o'l  mm.  The  cercaria 
lies  folded  in  the  cyst,  length  0*5  mm.  breadth  01  mm.  It  tapers 
posteriorly.  Skin  at  first  covered  with  fine  spines,  disappearing  as  they 
grow  older.     Body  dotted  with  fine  pigment. 

Theirs/  intermediate  host  is  still  unknown. 

Sub-family.     Metorchlinae,  Llihe,  1909. 
Genus.  Metorchis,  Looss,  1899,  emend,  auctor. 

Hind  end  rounded.  Gut  forks  reach  extreme  end.  Testes  only  slightly  lobed, 
filling  the  hind  end. 

Metorchis  truncatus,  Rud.,   18 19. 

This  species,  which  attains  a  length  of  2  mm.,  is  slender  and  conical, 
the  anterior  end  is  pointed  and  the  posterior  truncated,  and  provided 
with  a  muscular  tuberosity  that  resembles  a  terminal  sucker  ;  for  this 
reason  the  discoverer  of  the  species  (Rudolphi)  classed  it  with  the 
Amphistomes.  The  cuticle  in  the  young,  as  well  as  in  the  adult 
specimens,  is  entirely  and  closely  covered  with  spines.  Suckers  about 
equal  in  size  (0-134  to  0-172  mm.) ;  the  ventral  sucker  lies  somewhat 


262 


THE   ANIMAL   PARASITES   OF   MAN 


y.sc. 


in  front  of  the  middle  of  the  body.     The  phra-ynx  is  small  (0-09  mm.), 
the   oesophagus    minute,    the    intestinal  caeca  reach  to  the   posterior 

extremity.  Betw  een  them,  and  in  front  of 
their  blind  ends,  lie  the  two  elliptical  testes, 
one  generally  a  little  in  front  of  the  other. 
In  front  of  them,  either  in  the  median 
line  or  somewhat  laterally,  the  spheroidal 
ovary  is  situated ;  in  front,  again,  is  the 
uterus,  the  coils  of  which  usually  extend 
beyond  the  median  field.  The  vitellaria 
are  at  the  sides  of  the  central  third  of  the 
body,  thus  commencing  in  front  of  the 
ventral  sucker ;  cirrus  pouch  absent ;  the 
genital  pore  is  close  in  front  of  the 
acetabulum.  The  excretory  pore  is  ter- 
minal (?).     Eggs  29  /x  by  II  /JL. 

Metorchis  truncatus  lives  in  the  bile- 
ducts  of  the  seal,  cat,  dog,  fox,  and 
glutton  {Gulo  borealis).  The  source  of 
infection  is  unknown,  although  one  would 
suspect  fish.  Askanazy  did  not  succeed 
FiG.i62.—A/^jorckisjruncajus,     [^  getting  this  flukc  in  his  feeding  experi- 

Rud.  :  from  the  biliary  ducts  of  the  °  f  ^,  7#  ^        ,  • 

domestic  cat.    F.j.,  ventral  sucker ;     ments,    but    another    spccics,     Metorchts 
I.,  gut ;  v.sc.,  vitellaria ;  7".,  testes ;     alMdus,  not  uncommon  in  cats  by  feeding 

O.,  ovary ; -^.j.,  receptaculum  sem-        ,  '  i     /t  •  .1      x 

inis;  ^7/.,  uterus.   25/1.  them  on  roach  (Leuctscus  riitilus). 


R.s 


Family.     Heterophyiidae,  Odhner,  1914. 

Genus.     Heterophyes,  Cobbold,  1866. 

Syn.  :  Cotylogonimus,  Liihe,  1899;  Ccsnogommus,  Looss,  1899. 

No  crown  of  spines  on  head.  Body  divided  into  a  narrow,  movable,  anterior 
part  (neck),  and  a  broader,  less  movable,  posterior  portion,  which  contains  the 
genitalia.  The  suckers  separated  from  one  another  by  a  space  equal  to  half  the 
length  of  the  body  or  more ;  the  pharynx  is  close  behind  the  oral  sucker  ;  the 
oesophagus  is  long  ;  the  intestinal  caeca  extend  to  the  posterior  border  ;  the  genital 
pore  is  placed  laterally,  and  behind  the  ventral  sucker.  Genital  sucker  provided 
with  a  circlet  of  chitinous  rodlets,  shaped  like  stags'  horns.  The  testes  are  at  the 
posterior  end,  the  ovary  in  a  median  position  in  front  of  them.  Laurer's  canal  with 
receptaculum  seminis  present;  the  small  vitellaria  are  at  the  sides  of  the  posterior 
part  of  the  body.     Parasitic  in  the  intestine  of  mammals  and  birds. 


Heterophyes  heterophyes,  v.  Sieb.,  1852. 

Syn. :  Distotnmn  heterophyes,  v.  Siebold,  1852  ;  Heterophyes  cegyptica,  Cobbold^ 
1866;   Mesogoniinus  heterophyes,  Railliet,  1890;  Ccenogonimus  heterophyes,   Looss 
1900;  Cotylogonimus  heterophyes,  BrdMVi,  i<)oi. 


HETEROPHYES  HETEROPHYES 


263 


Length  up  to  2  mm.,  breadth  0*4  mm. ;  the  neck  not  sharply  defined  ; 
in  life  it  stretches  to  double  the  length  of  the  hind  body.  The  scales 
are  rectangular,  5  /z,  to  6  yu,  by  4  yLt,  their  posterior  margin  serrate  with 
seven  to  nine  teeth.  Cuticular  glands  are  numerous  on  the  ventral 
surface,  especially  in  the  fore  part  of  the  body,  and  partly  discharge 
at  the  anterior  border  of  the  oral  sucker.  The  oral  sucker  is  0*09  mm., 
the  ventral  sucker  o'23  mm.  in  diameter  ;  the  pharynx  measures 
0-05  to  0*07  mm.  in  length  ; 
the  oesophagus  is  about 
three  times  as  long  ;  pos- 
teriorly the  intestinal  caeca 
are  directed  one  towards 
the  other  and  terminate 
beside  the  excretory 
bladder.  Close  in  front 
of  the  posterior  ends  of 
the  intestinal  branches  are 
the  two  elliptical  testes, 
which  are  not  exactly  on 
the  same  level.  In  the 
middle  in  front  of  them  is 
the  receptaculum  seminis, 
and  in  front  of  the  latter 
lies  the  spherical  or  ellip- 
tical ovary.  The  two  vasa 
efferentia  unite  to  form  the 
vas  deferens,  which  after 
a  short  course  passes  over 
into  the  angularly  bent 
seminal  vesicle  ;  after  the 
entry  of  the  prostatic 
glands  it  becomes  united 
with  the  metraterm 
(vagina),  and  the  common 
duct  opens  into  the  genital 
sucker.  The  latter  is  some- 
what smaller  than  the  ven- 
tral sucker,  lateral  to  and 

close  (0*15  mm.)  behind  it,  and  bears  a  not  entirely  closed  ring  of 
from  seventy- five  to  eighty  chitinous  rods  (20  fx  in  length).  The  vitel- 
laria  on  either  side  consist  of  about  fourteen  acini.  The  uterus  is 
spread  almost  throughout  the  entire  posterior  part  of  the  body.  The 
eggs  have  thick  shells  with  a  knob  resembling  that  of  Clonorchis  eggs 
but  not  so  prominent,  and  measure  30  ^  by  17  ft;  they  contain  a 
completely  ciliated  miracidium  with  a  rudimentary  intestinal  sac. 


R.s. 


Fig.  163.  —  Heterophyes  helerophyesj  v.  Sieb. 
C.y  cerebral  ganglion;  /.,  intestinal  cseca ;  Ct.g., 
cuticular  glands  ;  V.sc,  viteljaria  ;  Ut.t  genital  sucker  ; 
7*.,  testes — the  excretory  bladder  between  them  ;  L.c, 
Laurer's  canal ;  R.s.^  receptaculum  seminis,  wiih  the 
ovary  in  front  of  it ;  G.c.y  ventral  sucker  ;  Vs.,  vesicula 
seminalis,  53/1.  On  the  left  side  above,  an  egg,  700/1, 
is  depicted,  and  below  it  three  chitinous  rodlets  from  the 
genital  sucker.     700/1.     (After  Looss.) 


?64 


THE   ANIMAL   PARASITES   OF   MAN 


This  species  was  discovered  in  185 1  by  Bilharz  in  the  intestine 
of  a  boy  who  died  in  Cairo  ;  a  second  case  was  only  found  in  1891 
and  published  by  R.  Blanchard,  so  that  it  appeared  as  if  the  species 
were  very  scarce.  According  to  Looss,  this  is,  however,  not  the  case, 
but  the  species  easily  escapes  notice  on  account  of  its  small  size. 
Looss  found  it  in  Alexandria  twice  in  nine  autopsies,  and  once  in 
Cairo,  and  has  recently  stated  that  in  man  *'  it  is  not  at  all  uncommon 
to  meet  with  the  parasite  in  cadavers,  and  the  eggs  of  the  worm  in 
the  stools  of  the  patients."  Leiper  records  one  case  from  Japan 
and  one  from  China.  The  parasites  occupy  the  middle  third  of  the 
small  intestine,  and  even  when  present  in  large  numbers  appear  to  be 

harmless. 

This  small  species,  according  to 
Looss,  frequently  occurs  in  Egyptian 
dogs,  less  so  in  cats,  and  has  also  been 
found  in  the  fox,  as  well  as  once  in 
M ilvus  parasiticus ;  Janson  also  reports 
this  species  from  the  intestine  of  the 
dog  in  Japan. 

Metagonimus,  Katsurada,  1913  ; 
Yokogawa,  Leiper,  1913. 

Resembles  in  general  structure 
Heterophyes.  In  the  arrangement  of 
its  ventral  genital  suckers  resembles 
but  differs  from  that  of  Tocotrema,^ 
Looss.  The  ventral  and  genital  suckers 
lie  laterally  and  on  the  right. 

Metagonimus  yokogawai.    Katsurada, 
1913. 

Syn. :    Yokogawa  yokogawai^  Leiper,  1913. 

One  to  1*5  mm.  long,  seldom  2-5  mm., 
and  0-4  to  07  mm.  broad  ;  elliptical  in 
shape.  The  body  is  thickly  covered 
with  nail-shaped  spines  about  10  /x 
long.  Oral  sucker  77  yu,  to  85  ji  in  diameter.  Ventral  sucker  charac- 
teristic and  peculiar  o"i2  to  0*14  mm.  by  o-o8  to  i  mm.  It  is  a  sac-like 
organ  placed  deeply  in  the  body,  but  does  not  open  as  in  other  flukes  on 
the  ventral  surface.  Testes  elliptical,  not  quite  symmetrically  placed  at 
the  hind  end  of  the  body.  Vesicula  seminalis  retort-shaped,  situated 
transversely,  internal  to  the  ventral  sucker.     Pars  prostatica  present. 


Fig.  164. — Metagonimus  yokogawai, 
Katsurada,  1913:  the  spines  are  only 
shown  over  a  small  patt  of  the  skin. 
(After  Leiper.) 


In  the  genus  Tocotrema  the  common  genital  duct  opens  into  the  ventral  sucker. 


DICROCCELIID^ 


265 


M.S. 


—    V.s. 


Ejaculatoiy  duct  opens  with  the  uterus  into  a  genital  sinus,  which, 
together  with  the  internal  opening  of  the  ventral  sucker,  opens  into  a 
pit  at  the  front  of  the  ventral  sucker.  The  opening  of  the  genital  sinus 
and  that  of  the  ventral  sucker  are  furnished  with  a  complex  muscular  ap- 
paratus. Ovary  spherical,  0-12  to  0*13  mm.  in  diameter,  lies  in  the  middle 
of  the  hind  body.  Receptaculum  seminis  and  Laurer's  canal  present. 
Vitellaria  in  the  hind  half  of  the  body,  consisting  of  about  ten  acini  on 
each  side.  Shell  gland  to  the  left  of  the 
ovary.  Uterus  forms  three  to  four  trans- 
verse coils.  Eggs  elliptical,  double 
contoured,  yellowish-brown  in  colour. 
There  is  no  shoulder  below  the  oper- 
culum as  in  the  eggs  of  CI.  sinensis. 
At  the  rounder  end  there  is  a  thicken- 
ing or  knob  different  from  the  spine- 
like or  hook-like  process  seen  in  CI. 
sinensis.     Dimensions  28  //,  by  16  //,. 

Habitat.  —  Mainly  in  upper  or 
middle  portion  of  jejunum,  rarely  in 
caecum.  They  penetrate  deep  into  the 
mucosa,  but  not  into  the  submucosa, 
and  post  mortem  appear  as  a  number 
of  small  brown  points.  They  fre- 
quently occur  in  the  solitary  glands, 
which  they  destroy.  They  cause 
chronic  catarrh  of  the  gut.  Parasitic 
in  man  and  mammals. 

Geographical  Distribution. — Japan. 

Life- history. — The  cercarial  stage 
occurs  in  a  trout  {Plecoglossns  altivelis) 
and  seldom  in  Crassius  sp.  and  Cypri- 
nus  sp.  Infection  takes  place  through 
the  eating  of  the  fish  raw.  Seven  to 
sixteen  days  later  eggs  appear  in  the 
faeces  (of  dog). 

Family.   DIcrocoeliidae,  Odhner,  1910. 
Genus.     DicroccBlium,  Dujardin. 


V.SC, 


Fig.  165.  —  Dicrocoeluim  dendriticumy 
Rud.  V.s.y  ventral  sucker  ;  Cb.,  cirrus 
pouch;  /.,  intestinal  caeca;  K^^.,  vitel- 
laria ;  Z".,  testicles;  O.,  ovary;  M.s.^ 
oral  sucker  ;   ^A,  uterus.     15/1. 


DicrocoeliidcE.,  with  leaf-shaped  bodies, 
pointed  posteriorly  and  anteriorly.  Greatest 
width  behind  the  mid-line.     Vitellaria  double. 

The  testes  smooth  or  indented,  lying  symmetrically  or  obliquely  beside  or  behind 
the  ventral  sucker.  The  ovary  approaches  the  median  line  behind  one  testis. 
Parasitic  in  the  liver  and  gall-bladder  (rarely  in  the  intestine)  of  members  of  all 
classes  of  vertebrate  animals — by  preference  in  birds  and  mammals. 


266 


THE   ANIMAL   PARASITES   OF   MAN 


Dicrocoelium  dendriticum,  Rud.,  1819. 

Syn.  :  Dicroccelimn  lanceatum,  Stil.  and  Hass.,  1896  ;  Fasciola  lanceolata, 
Rud.,  1803  {nee  Schrank,  1790)  ;  Distomutn  lanceolatum,  Mehlis,  1825  ;  Dicrocoelium 
lanceolatum^  Dujardin,   1845. 

Body  lancet-shaped,  narrowing  especially  at  the  anterior  extremity  ; 
length  8  to  10  mm.,  breadth  I'S  to  2*5  mm.,  the  greatest  breadth  usually 
behind  the  middle  of  the  body.  Suckers  distant  from  each  other  by 
about  one-fifth  the  length  of  the  body ;  oral  sucker  about  o*  5  mm.,  ventral 
sucker  about  0*6  mm.  Pharynx  globular,  adjoining  the  oral  sucker; 
oesophagus  o-6  mm.  in  length  ;  intestinal  caeca  reach  to  four-fifths  of 
the  body  length.  Genital  pore  at  the  level  of  the  bifurcation  of  the 
intestine  ;  cirrus  pouch  small  and  slender.  The  large,  slightly  lobed 
testes  lie  obliquely  one  behind  the  other  behind  the  ventral  sucker ; 


Fig.  166. — Eggs  of  Dicro- 
cceliu77i  dendriticum^  Rud. 
To  the  left  seen  flat,  to  right 
lying  on  one  side.     600/1. 


Fig.  167.  —  Miracidia 
Dicrocoelium  dendriticum.  a, 
from  the  dorsum  ;  d,  from  the 
side.     (After  Leuckart.) 


the  ovary,  which  is  considerably  smaller,  is  placed  behind  the  pos- 
terior one  ;  the  vitellaria,  commencing  at  the  level  of  the  posterior 
testis,  terminate  far  before  the  caeca.  The  uterus,  situated  behind  the 
ovary,  extends  throughout  the  posterior  end,  not  confined  to  the  central 
field,  but  overlapping  the  lateral  fields  with  its  transverse  coils ;  at 
the  posterior  edge  of  the  body  it  turns  back  again  and  winds  forwards 
to  the  ovary  in  transverse  loops,  then  between  the  testes,  and  finally, 
dorsal  to  the  ventral  sucker,  terminates  in  the  genital  pore.  The 
thick-shelled  eggs  when  young  are  yellowish,  when  older  dark  brown. 
They  measure  38  /x  to  45  yL6  by  22  //,  to  30  /n.  They  contain  an  oval  or 
roundish  miracidium,  only  the  anterior  part  of  which  is  ciliated,  and 
which  possesses  a  rudimentary  intestinal  sac  with  a  boring  spine. 
The  miracidia  do  not  hatch  out  in  water  spontaneously,  but,  accord- 
ing to  Leuckart,  in  the  intestines  of  slugs  {Limax,  Arion),  but  they 
do  not  develop  either  in  these  (slugs)  or  in  water-snails. 

The  lancet  fluke  inhabits  the  biliary  duct  of  herbivorous  and 
omnivorous  mammals  (sheep,  ox,  goat,  ass,  horse,  deer,  hare,  rabbit, 
pig),  and  is  often  found  associated  with  the  liver  fluke  ;  it  is  not, 
however,  so  common  nor  so  widely  disseminated,  nevertheless,  it  has 
been  met  with  outside  of  Europe,  namely,  in  Algeria,  Egypt,  Siberia, 
Turkestan,  and  North  and  South  America. 


ECHINOSTOMID^.  267 

In  man  it  is  still  more  uncommon  than  the  liver  fluke,  and  has 
hitherto  only  been  observed  seven  times  (Germany,  Bohemia,  Italy, 
France,  and  Egypt) ;  it  may,  however,  have  occurred  more  frequently, 
and  have  been  overlooked,  as  in  slight  infections  it  produces  no 
special  symptoms. 

The  intermediate  host  is  still  unknown.  Leuckart  for  some  time 
held  the  opinion  that  small  species  of  Plaiiorbis  from  fresh  water, 
which  contain  encysted  Distomata,  were  to  blame,  and  he  supported 
his  views  by  a  feeding  experiment  which  seemingly  yielded  positive 
results;  this,  however,  is  not  definitely  proved.  Piana's  statement 
that  small  land  snails  are  the  intermediate  hosts  has  also  not  been 
proved. 

Family.     Echinostomidae,  Looss,  1902. 

Sub-family.     Echinostomlnae,  Looss,  1899. 

Genus.     Echinostoma,  Rud.  1809  )  Dietz,  1910. 

Fore-body  not  bulging.  Greatest  width  at  or  behind  the  ventral 
sucker.  Oral  sucker  not  atrophied.  Collar  kidney-shaped  with  a  double 
dorsally  unbroken  row  of  spines,  terminating  in  four  to  five  angle  spines. 
The  border  spines  of  the  aboral  series  not  larger  than  the  oral.  Skin 
spined  or  smooth.  Body  elongated.  Uterus  long  with  numerous 
transverse  coils.  Ventral  sucker  in  the  anterior  quarter  of  body.  Cirrus 
sac  small,  almost  completely  in  front  of  the  ventral  sucker.  Testes  round 
or  oval,  smooth  incurved  or  lobed,  in  the  hinder  half  of  body.  Ovary 
median  or  lateral  in  front  of  testes.  Vitellaria  from  hinder  margin  of 
ventral  sucker  to  end  of  body.    Eggs  oval,  84  /x  to  126/^  by  48  jjl  to  82  /i,. 

The  spines  placed  most  ventrally,  or  those  placed  most  medially 
on  ventral  surface,  are  from  difi"erences  of  position  or  form  termed 
"  angle  "  spines,  the  rest  "  border  "  spines. 

Type. — Echinostoma  echinatum,  Rud. 

Echinostonna  ilocanum,  Garrison,  1908. 

Length  4  to  5  mm.,  breadth  i  to  1-35  mm.,  thickness  0-5  to 
0-6  mm.  The  circum-oral  disc  o"3  mm.  broad,  separated  by  a  shallow 
groove  from  the  body.  Crown  of  forty-nine  spines  and  five  to  six 
angle  spines  on  each  side  continuous  with  an  irregularly  alternat- 
ing series  of  fourteen  spines  on  the  dorsum.  Largest  spines  are 
34  fi  long,  8  /JL  thick  at  the  base.  The  remainder  of  the  dorsal  spines 
are  24  yLtby  6  yit.  Skin  thickly  covered  with  scales  on  the  margins 
of  the  body  as  far  back  as  the  level  of  the  hind  testis.  Oral  sucker, 
o-i8  mm. ;  ventral  sucker,  0*4  to  0*46  mm.  Its  anterior  border  about 
Q-oy  mm.  from  the  anterior  end.  Pharynx  0*17  mm.  long,  cii  mm. 
broad.  Testes  about  mid-line  of  the  body,  much  lobed ;  the  lobes  of 
the  anterior  testis  run  transversely,  while  the  axis  of  the  posterior  testis 
is   longitudinal,  as  often   occurs    in    the   Echinostomidce.     Cirrus  sac 


268 


THE   ANIMAL   PARASITES   OF   MAN 


..-Vo. 


reaches  to  the  centre  of  the  ventral  sucker.  Ovary  transversely  oval 
in  front  of  the  testes.  Vitellaria  commence  about  half-way  between 
the  ventral  sucker  and  ovary  and  extend  to  the  posterior  end.  Eggs 
numerous,  92  /x  to  ii4/>«'  by  53  //,  to  82 /a. 

Average.— gg'SH'hy  56/^. 
Habitat. — Gut  of  man  (Filipinos), 
Philippine  Islands. 

Echlnostoma  malayanum, 

Leiper,   191 1. 

Twelve  millimetres  long,  3  mm. 
broad,  1*3  mm.  thick.  Ends  bluntly 
rounded.  At  the  anterior  end  a 
ventral  furrow  on  either  side,  one- 
third  the  width  of  the  body, 
marking  off  the  circum-oral  collar. 
Along  its  edge  is  a  row  of  forty- 
three  spines  extending  across  the 
middle  line  dorsally  but  not  ven- 
trally.     The  spines  vary  in  size  from 


Fig.  168.  —Echinostoma  ilocanum.  Vo.^ 
oral  sucker ;  Ph.^  pharynx  ;  Cirre,  cirrus  ; 
F.z;.,  ventral  sucker;  6V.,  uterus;  G.c.^ 
ovary;  Ov.,  shell  gland  ;  T.,  testes; 
7".^.,  vitellarium  ;  C.ex,,  excretory  vesicle. 
(After  Erumpt.) 


Fig.  169.  —  Echinostoma  ilocanum^ 
Garrison,  1908  :  head  end  showing  collar 
of  spines,  ventral  view.     (After  Leiper.) 


0*07  mm.  in  length  (ventrally)  to  0*05  to  o'oi6  mm.  (dorsally).  Cuticular 
spines  also  exist  on  the  ventral  side  as  far  back  as  posterior  end  of 
body,  but  dorsally  limited  to  a  triangular  area  ending  in  front  of  the 
ventral  sucker.     Oral  sucker  0*07  mm.  thick,  occupying  the  middle 


HIMASTHLIN.E 


2fy) 


third  of  the  circum-oral  disc;  pharynx  0*25  mm.  in  diameter;  oeso- 
phagus 0-04  mm.  long ;  gut  caeca  simple,  extending  to  end  of  body ; 
ventral  sucker  0*9  mm.  long  by  075  mm.  broad  by  07  mm.  deep  ;  wall 
about  0*25  mm.  thick.  The 
sucker  is  inclined  at  an  angle 
of  40°  to  the  ventral  surface. 
Testes  lobed,  one  behind  the 
other,  behind  the  ventral  sucker. 
Cirrus  pouch  well  developed, 
reaching  to  the  posterior  edge 
of  the  sucker.  Genital  pore  in 
the  angle  between  neck  and 
anterior  lip  of  ventral  sucker. 
Ovary  smooth,  0-3  mm.  in 
diameter,  0*85  mm.  behind 
ventral  sucker.     Vitellaria  very 

numerous,  extending  from  posterior  margin  of  sucker  to  posterior  end 
of  body,  where  they  intermingle.  Eggs  few  in  numb  :r,  brown  and  large. 
Habitat. — Gut  of  man  (Tamils),  Malay  States. 


Fig.  170. — Echinostoma  malayanum,  Leiper, 
1912  :    anterior    end  showing  collar  of  spines, 
(After  Leiper.) 


ventral  view. 


Sub-family.     Himasthlinae,  Odhner,   1910. 

Genus.     Artyfechinostomum,  Clayton-Lane,  1915. 

Crown  of  thirty-nine  spines,  continuous  over  dorsum.  Two  corner 
spines  long.  Vitellaria  extend  from  posterior  margin  of  sucker  to  pos- 
terior end  of  fluke.  Eggs  without  filament.  [Although  the  possession 
of  strong  rose-thorn  hooks  is  given  by  Odhner  as  a  sub-family  charac- 
teristic, yet  in  this  genus  assigned  to  this  sub-family  they  have  not 
been  seen.— J.  W.  W.  S.] 

Artyfechinostomum  sufrartyfex,  Clayton-Lane,  1915. 

Spirit  specimens  :  9  by  2*5  by  o'8  mm.  thick.  Ventral  sucker 
conspicuous,  i  mm.  in  diameter.  Cirrus  sac  2  mm.  long.  Testes 
lobed,  about  1*5  mm.  in  diameter.  Posterior  border  of  posterior 
testes  I  mm.  from  posterior  end.  Vitellaria  meet  posteriorly  behind 
the  posterior  testis. 

Family.     Schistosomidae,  Looss,  1899. 
Genus.     Schistosoma,  Weinl,  1858. 

Syn. :  GyncEcophorus,  Dies.,  1858;    Bilharzia,  Cohh.,  1859;    Thecosoma^ 

Moq.  Tandon,   i860. 

The  males  have  bodies  that  widen  out  considerably  behind  the  ventral  sucker, 

the  lateral  parts  of  which  in-roll  ventrally,  forming  the  almost   completely  closed 

canalis   gynsecophorus,  within  which  the  female  is  enclosed.      There  is  no  cirrus 

pouch.      The  male  has  five  or  six  testes,  the  females  are  filiform  ;  the  uterus  is  long. 


270 


THE   ANIMAL    PARASITES   OF   MAN 


There  is  no  Laurer's  canal.  The  ova  almost  equally  attenuated  at  either  extremity  ; 
they  have  a  small  terminal  spine,  and  are  not  provided  with  a  lid.  They  contain 
a  miracidium,  ciliated  on  all  sides,  which  is  characterized  by  the  possession  of  two 
large  glandular  cells,  which  discharge  anteriorly  beside  the.  gastric  sac.  They  live 
in  the  vascular  system  of  mammals.  (An  allied  genus  [Bilharziella]  lives  in  the 
blood-vessels  of  birds.) 

Schistosoma  haematobium,   Bilharz,  1852. 
Syn. :   Distoma  hceynatobiujn^  Bilh. ;   Distoma  capense^  Harley,  1864. 

The  Male  is  whitish,  12  to  14  mm.  in  length,  but  is  already  mature 
when  4  mm.  long.  The  anterior  end  is  o*6  mm.  or  a  little  over  in 
length.  The  suckers  are  near 
each  other,  the  oral  sucker  is  in- 
fundibular, and  the  dorsal  lip  is 
longer  than  the  ventral  one.  The 
ventral  sucker  is  a  little  larger, 
0*28  mm.,  and  is  pedunculated. 
A  little  behind  the  ventral  sucker 
the  body  broadens  to  a  width  of 
I  mm.,  decreasing,  however,  in 
thickness  ;  the  lateral  edges  in-roll 
ventrally,  so  that  the  posterior 
part  of  the  body  appears  almost 
cylindrical,  0*4  to  0*5  mm.  in 
diameter  ;  the  posterior  extremity 
is  somewhat  more  attenuated. 
The  dorsal  surface  of  the  pos- 
terior part  of  the  body  is  covered 
with  spinous  papillae.  There  are 
delicate  spines  on  the  suckers,  and 
largerones  invest  the  entire  internal 
surface  of  the  gynaecophoric  canal, 
as  well  as  a  longitudinal  zone  at 
the  edge  of  that  side  of  the  external 
surface  that  is  covered  by  the  other 
side  rolling  over  it.  The  oeso- 
phagus is  beset  with  numerous 
glandular  cells  (fig.  173),  and  pre- 
sents two  dilatations;  the  intestinal 
bifurcation  is  close  in  front  of 
the  ventral  sucker,  the  two 
branches  uniting  sooner  or  later  behind  the  testes  into  a  median 
trunk,  which  may  again  divide  at  short  intervals.  The  excretory 
pore  is  at  the  posterior  end,  but  placed  somewhat  dorsally  ;  the 
genital  pore  is  at  the  beginning    of    the    gynaecophoric    canal,  thus 


Fig.  \T\.— Schistosoma  hcejftatobtum,B\\.  : 
male  carrying  the  female  in  the  canalis  gynge- 
cophorus.     1 2/ 1.     (After  Looss.) 


SCHISTOSOMA   H^MATOBIUM 


271 


Fig.  172.  —  Tiansverse  section 
through  a  pair  of  Schistosoma 
hcematobium  in  copula.  In  the  male 
the  p6int  of  reunion  of  the  intestinal 
forks  has  been  cut  across.  (After 
Leuckart.) 


O.s. 


behind  the  venti*al  sucker  ;  into  it  opens  the  vas  deferens  which, 
posteriorly,  broadens  into  the  seminal  vesicle  and  then  continues 
as  the  vasa  efferentia  of  the  four  or  five  testes  (fig.  173). 

The  Female — filiform,  about  20  mm.  in  length,  pointed  at  each  end, 
and  measuring  0*25  mm.  in   diameter  in 
the  middle.     Their  colour  varies  accord- 
ing to  the  condition   of  the  contents  of 
the  intestine.     (Posteriorly  they  are  dark 
brown  or  blackish.)   The  cuticle  is  smooth 
except  in  the  sucker,  where  there  are  very 
delicate  spines,  and  at  the  posterior  end, 
where  there  are  other  larger  spines.     The 
oral    sucker    is   a   little    larger    than   the 
pedunculated    ventral    sucker    (0*07    and 
0*059    mm.    respectively).      The    anterior 
part  of  the  body,  0*2  to  0*3  mm.  in  length  ; 
the  oesophagus    is  as  in  the  male.     The 
intestinal   bifurcation  is  in   front   of   the 
ventral  sucker,  the  two  branches  uniting 
behind  the  ovary  and  the  trunk  running 
in  a  zigzag  manner  to  the  posterior  border. 
There  are  indications  of  diverticula  at  the 
flexures.    The  ovary  is  median.    In  young 
females  it  is  of  an  elongated  oval  shape ; 
in  older  females  the  posterior  end  becomes 
club-shaped,    whereas    the   anterior    end 
becomes  attenuated  ;  the  oviduct  origin- 
ates  at  the   posterior  end,  but    immedi- 
ately turns  forwards  and  joins  the  parallel 
vitelline  duct  in  front  of  the  ovary  (fig.  174), 
where   the   shell   gland  cells   open  ;    the 
common   canal  becomes  dilated  to  form 
the    ootype,    and   then    proceeds    as   the 
uterus,    with     only    slight    convolutions, 
along  the  central  field  to  the  genital  pore, 
which  lies  in  the  middle  line  immediately 
behind   the    ventral    sucker.     The   single 
vitellarium  starts   behind   the   ovary   and 
extends  to  the  posterior  end.     The  acini 
are  situated  at  the  sides  of  the  excretory 

duct,  which  runs  a  median  course.  The  eggs  are  compact  spindles, 
much  dilated  in  the  middle  ;  they  have  no  lid,  and  are  provided  with  a 
terminal  spine  (rudimentary  filament)  at  the  posterior  end,  measuring 
120  yu,  to  150  /I,  in  length  and  40  /^  to  60  /a  in  breadth,  but  vary  in  size 
and  shape  (fig.  175;. 


Oe.  _ 


V.s.~- 


Fig.  173. — Anterior  end  of  the 
male  Schistosoma  hcematobium,  Bilh. 
V,s.,  ventral  sucker  ;  /.,  gut  cceca  ; 
G.p.,  genital  pore  ;  T.,  testes  ; 
O.s.y  oral  sucker  ;  Oe.,  cesophagus 
with  glandular  cells;  V.s.,  vesicula 
seminalis.     40/1.     (After  Looss.) 


272  THE   ANIMAL   PARASITES   OF   MAN 

Distribution. — In  order  to  understand  the  distribution  of  the  worms 
and  eggs  in  the  body,  it  may  be  well  to  recall  the  blood  supply  of  the 
abdominal  and  pelvic  organs.  It  is  generally  assumed  that  the  early 
life  (?cercarial  stage)  of  the  worms  occurs  in  the  liver,  and  that  the 
young  worms  travel  from  here,  where  they  are  invariably  found,  to 
their  various  sites  along  the  portal  vein  and  its  tributaries  and  so 
against  the  blood  stream.     The  tributaries  of  the  portal  vein  are  : — 

(i)  Superior  mesenteric,  the  tributaries  of  which  are  :  (a)  the  veins 
of  the  small  intestine  ;  (6)  ileo-colic  ;  (c)  right  colic ;  (d)  middle  colic  ; 
(e)  right  gastro-epiploic  ;  and  (/)  inferior  pancreatic.  By  these  paths 
infection  of  the  small  intestine,  ascending  and  transverse  colon  and 
pancreas  would  occur. 

(2)  Splenic.     (Ova  have  been  recorded  by  Symmers  in  the  spleen.) 

(3)  Inferior  mesenteric,  the  tributaries  of  which  are  (a)  superior 
haemorrhoidal  veins  from  the  upper  part  of  the  haemorrhoidal  plexus; 
(6)  sigmoid  veins  from  sigmoid  flexure  and  lower  portion  of  descending 
colon  ;  (c)  left  colic  vein  draining  descending  colon. 

The  superior  haemorrhoidal  veins  form  a  rich  plexus  in  the  rectum, 
and  below  this  level  in  the  upper  and  middle  parts  of  the  anal  canal. 
The  plexus  forms  two  networks,  an  internal  plexus  in  the  submucosa 
and  an  external  on  the  outer  surface.  The  internal  plexus  opens  at 
the  anal  orifice  into  :  (a)  branches  of  the  inferior  haemorrhoidal  vein 
(from  the  pudic)  ;  (b)  the  external  plexus.  The  external  plexus  gives 
off :  (a)  inferior  haemorrhoidal  opening  into  internal  pudic  (of  internal 
iliac  vein) ;  (b)  mid-haemorrhoidal  into  internal  iliac  or  its  branches  ; 
and  (c)  superior  haemorrhoidal  opening  into  inferior  mesenteric. 
The  external  plexus  further  communicates  with  the  vesico-prostatic 
plexus.  The  vesico-prostatic  (vaginal)  plexus  opens  into  the  vesical 
veins,  which  drain  into  the  interior  iliac  vein.  This  plexus  also 
receives  afferents  from  the  pudendal  plexus,  the  chief  tributary  of 
which  is  the  dorsal  vein  of  the  penis.  The  pudendal  plexus  also 
receives  branches  from  the  inferior  pudic  and  the  anterior  surface  of 
the  bladder. 

There  is  thus  a  communication  between  the  portal  vein  and  the 
vena  cava  by  means  of  these  plexuses,  viz.,  through  the  inferior  and 
middle  haemorrhoidals,  and  by  the  inferior  haemorrhoidals  to  the 
bladder  and  thence  by  the  vesical  veins  or  the  pudic  to  the  caval 
system  (interior  iliac). 

It  is  thus  by  the  inferior  mesenteric  and  its  tributaries  that  the 
worms  reach  the  descending  colon,  rectum,  anal  canal,  and  eventually 
the  bladder,  and  in  some  cases  the  caval  system. 

Before  considering  what  is  actually  found  post  mortem  in  these 
veins  and  the  organs  drained  by  them,  we  may  further  recall  the  fact 
that  the  calibre  of  ''medium"  veins  is  4  to  8  mm.,   ''small"  veins 


SCHISTOSOMA   H^MATOBIUM  273 

less  than  40//,  in  diameter  and  capillaries  Syu,  to  20 /x.  Further,  the 
maximum  diameter  of  the  male  worm  is  1  mm.,  that  of  the  female 
2S0  fjL  and  eggs  ///  uiero  So  fi  to  90//,  long  by  30//,  to  40//,. 

Liver  and  Portal  Vein. — Here  worms  are  most  easily  found  post 
mortem.  Often  only  males  are  found  and  these  of  the  same  size, 
and  if  females  occur  only  a  few  worms  are  found  in  copula.  The 
worms  are  frequently  not  full  size  and  the  males  may  contain  no  free 
spermatozoa  in  their  testes,  and  as  regards  the  females  some  may 
be  fertilized,  others  not,  as  shown  by  the  presence  or  absence  of 
spermatozoa  in  the  seminal  receptacle  or  uterus.  In  either  case  they 
may  contain  eggs — lateral-spined — usually  one,  less  often  two,  but 
there  may  be  as  many  as  five  or  six.  These  eggs  may  also  show 
some  abnormality,  which  takes  the  form  of :  (i)  abnormal  con- 
tents, viz.,  disintegrating  yolk  cells  with  or  without  an  ovarian  cell  ; 
(2)  abnormal  shape  but  with  normal  contents  and  probably  repre- 
sented by  the  collapsed  and  empty  egg-shells  which  are  found  in  the 
tissues. 

As  to  the  interpretation  of  these  facts,  Looss  believes  that  these 
lateral-spined  eggs  are  products  of  young  females  whose  egg-laying 
is  not  at  first  properly  regulated.  The  shape  that  the  eggs  take,  viz.y 
with  a  lateral  spine,  is  determined  by  an  excess  of  material — ovarian 
and  yolk  cells — being  present  in  the  ootype.  The  shape  of  eggs 
depends  upon  the  position  they  have  in  the  ootype  during  their 
formation.  In  young  females  an  excess  of  cells — yolk  cells  especially 
— accumulates,  distending  not  only  the  dorsal  wall  but  a  portion  also 
of  the  short  duct  joining  the  ootype  to  the  uterus.  The  result  of  this 
is  that  the  axis  of  the  ootype  and  egg  is  almost  transverse  to  the  body, 
and  the  posterior  funnel-shaped  portion  of  the  ootype,  instead  of 
being  terminal,  has  now  a  lateral  or  rather  a  ventral  position,  so  that 
the  spine  which  occupies  this  portion,  instead  of  being  terminal,  is 
now  lateral.  It  is  noteworthy  that  these  lateral-spined  eggs  are 
thicker,  owing  to  the  excess  of  material  present,  and  not  uncommonly 
have  a  curved  anterior  border,  due  to  a  projection  of  the  anterior  end 
into  the  anterior  opening  of  the  ootype. 

As  these  eggs  are  being  laid  by  females  in  the  portal  vein  they  are 
carried  back  to  the  liver  by  the  blood  stream.  The  liver  is  one  of 
the  commonest  sites  for  these  eggs ;  also  terminal-spined  eggs  may  be 
found  here  for  the  same  reason. 

Hcemorrhoidal  Veins. — Mature  worms,  generally  in  copula,  are 
usually  found  here,  though  young  not  fully  grown  females  may  also 
occur.  The  tissues  of  the  rectal  wall  (or  colon)  show,  as  a  rule,  large 
quantities  of  lateral-spined  eggs,  though  less  often  only  terminal- 
spined  eggs  may  be  found. 

Vesico-prostatic  Plexus. — Worms  in  copula  are  found  in  the  veins 
18 


274  THE   ANIMAL   PARASITES   OF   MAN 

of  the  submucosa  in  the  bladder,  and  the  eggs  in  the  mucosa,  and 
those  voided  are  usually  terminal-spined,  though  lateral-spined  eggs 
are  not  so  rare  as  generally  thought.  The  problem  next  arises  as  to 
how  the  eggs  get  to  the  lumen  of  the  gut  or  bladder. 

The  female  worm  is  280  /x  in  diameter.  Veins  in  the  submucosa 
of  the  rectum  less  than  178 /x  in  diameter  are  not  affected  with  endo- 
phlebitis.  It  is  probable  that  the  female  even  by  stretching  could  not 
penetrate  much  beyond  this.  Eggs  are  probably  then  laid  in  the 
submucosa  as  near  the  muscularis  mucosa  as  possible.  Now  if  the 
eggs  are  laid  in  a  vein  of  larger  calibre  than  the  worm  fills,  the  eggs 
would  be  carried  back  to  the  inferior  mesenteric  vein,  so  that  pre- 
sumably the  worm  must  succeed  in  blocking  the  vein  already  narrowed 
by  endophlebitis,  so  that  by  the  stasis  which  ensues  the  eggs  may 
escape  from  the  veins.  How  this  occurs  is  not  exactly  known  ;  it  is 
not  necessarily  due  to  the  spine,  as  the  same  escape  into  the  tissues 
occurs  in  spineless  eggs,  such  as  those  of  Schistosoma  japonicum.  The 
eggs,  then,  pass  as  foreign  bodies  through  the  tissues.  Another  hypo- 
thesis is  that  the  worms  leave  the  veins  in  order  to  lay  their  eggs, 
but  the  evidence  is  against  this. 

Caval  System. — Occasionally  worms  that  have  passed  through  the 
vesical  plexus  may  be  found  in  the  iliac  vein,  inferior  vena  cava, 
and  even  the  lungs.  If  the  worms  are  young  they  contain  a  lateral- 
spined  €^gg ;  if  adult,  numerous  (50  to  100)  terminal-spined  eggs. 

Lungs. — When  the  liver  is  strongly  infected  with  (terminal-spined) 
eggs  it  is  possible  that  by  passive  movements  some  may  pass  into  the 
intralobular  veins,  and  thence  by  the  inferior  vena  cava  to  the  lungs. 

Gall-bladder. — Similarly  terminal-spined  eggs  pass  into  the  bile- 
capillaries  and  gall-bladder  (where  they  may  be  abundant),  and  so  into 
the  faeces. 

Detection  of  Eggs. — Occasionally  eggs  may  be  found  in  various  other 
parts  of  the  body.  They  are  best  detected  by  macerating  pieces  of  the 
tissue  in  question  in  about  ^  per  cent,  hydrochloric  acid  at  50  to  60°  C. 
(Looss). 

Pathological  changes : — 

Rectum.— These  have  been  studied  thoroughly  by  LetuUe  in  the 
case  of  an  apparently  pure  infection  of  the  rectum.^  They  take  the 
form  of  a  chronic  diffuse  inflammation,  which  may  result  in— (i)  ulcera- 
tion, or  (2)  hyperplasia  of  the  mucosa,  producing  adenomata. 

Ulcerative  Form. — The  mucosa  is  transformed  into  a  mass  of 
vascular  connective  tissue.  The  connective  tissue  spaces  next  be- 
come invaded    by  numerous   mononuclear  cells.      The   tissue    itself 


'  It  is  noteworthy  that  in  this  almost  classical  case  no  worms  were  found  in  any  of  the 
sections.  It  is  further  noteworthy  that  the  eggs  in  the  rectum  showe-i  great  irregularity  of 
form.  Eggs  with  a  spine  at  each  end  were  not  uncommon  ;  exceptionally  eggs  with  two 
polar  spines  and  one  lateral. 


SCHISTOSOMA   H^MATOBIUM 


275 


undergoes  diffuse  sclerosis,  becoming  hard  and  fibroid.  Eventually 
ulcerative  necrosis  sets  in.  During  these  changes  the  Lieberkiihn 
glands  are  destroyed.  The  process  does  not  extend  to  the  submucosa, 
in  this  respect  differing  from  that  in  chronic  dysentery. 

Hyperplastic  Form. — The  Lieberkiihn  glands  of  the  mucosa  at  first 
hypertrophy  ;  then  there  is  an  actual  hyperplasia  resulting  in  adeno- 
mata. The  interstitial  tissue  of  the  glands  is  also  greatly  hyper- 
trophied,  giving  rise  to  very  vascular  granulations.  These  growths 
are  often  hollow  and  contain  worms.  Many  eggs  are  found  in  the 
mucosa  on  their  way  to  the  lumen  of  the  gut. 

The  muscularis  mucosa  is  thickened  up  to  twice  or  even  ten  times 
the  normal.  Its  vessels  are  dilated  (36  fi  to  80  /i,),  but  they  do  not 
allow  of  the  passage  of  worms. 

The  submucosa  is  profoundly  changed ;  rigid  and  hard  instead  of 
supple.  It  is  here  that  the  greatest  number  of  eggs  occur.  A  re- 
markable condition  of  endophlebitis  exists  in  the  veins  of  the 
submucosa,  not  only  in  the  smaller  ones  but  also  in  the  larger  ones 
(370//,  by  270 /x).  This  endophlebitis  results  in  a  more  or  less  complete 
occlusion  of  the  vessels  of  the  lumen. 

The  muscular  coats  are  free  from  change,  also  their  veins. 

The  Serous  Coats. — The  veins  about  1,900//,  also  show  endophlebitis. 
Besides  the  rectum,  in  extreme  cases  even  the  transverse  colon,  the 
caecum  and  small  intestine  may  be  affected. 

Bladder. — In  the  early  stages  the  mucosa  is  deep  red  and  swollen 
like  velvet,  or  there  may  be  localized  patches  of  hyperaemia  or 
extravasation.     The  subsequent  changes  take  two  chief  forms  : — 

(i)  Sandy  Patches. — The  mucosa  ooks  as  if  it  were  impregnated 
with  a  fine  brownish  or  yellowish  powder  (myriads  of  ova).  This  is 
accompanied  by  a  gradual  hypertrophy  and  new  formation  of  con- 
nective tissue,  so  that  dry,  hard  or  plate-like  patches  with  this  sandy 
appearance  arise ;  the  thickening  eventually  affects  all  the  coats  of 
the  bladder.  In  the  older  patches  many  of  the  eggs  are  calcified. 
These  patches  sooner  or  later  break  down,  ulcerate  and  necrose. 
Phosphatic  deposits  are  abundant  and  stone  is  common.  These 
patches  are  not  found  in  the  rectum. 

(2)  Papillomata. — Where  the  inflammatory  change  produced  by 
the  eggs  gives  rise  to  hypertrophy  and  hyperplasia  of  the  mucosa, 
papillomata  result,  the  axis  of  which  is  formed  by  connective  tissue 
of  the  submucosa.  These  are  most  variable  in  shape  and  form  and 
bleed  readily,  and  sometimes  contain  cavities  of  extravasated  blood. 

As  in  the  rectum,  it  is  in  the  submucosa  that  eggs  are  most 
abundant,  and  worms  in  copula  occur  in  the  veins  of  this  layer,  but 
endophlebitis  is  not  as  general  as  described  in  the  rectum.  Malignant 
disease  of  the  bladder  is  not  an  uncommon  sequela  of  bilharziasis. 


276 


THE   ANIMAL   PARASITES   OF   MAN 


Besides  the  bladder,  the  ureters  and  kidneys  may  in  advanced  cases 
be  involved.  The  prostate  and  vesiciilae  seminales  are  commonly 
diseased.  Eggs  have  been  recorded  in  the  semen.  The  urethra  is 
frequently  attacked;  the  vagina  in  the  female. 

Eggs  also  occur  in  the  lymphatic  glands  of  the  gut. 

Geographical  Distribution.  —  East  Africa  : 
Nile  Valley,  Red  Sea  Coast,  Zanzibar,  Portu- 
guese East  Africa,  Delagoa  Bay,  Natal,  Port 
Elizabeth. 

South  Africa  :  Cape  Colony,  Orange  Free 
State,  Transvaal,  Mauritius,  Bourbon,  Mada- 
gascar. 

West  Africa  :  Angola,  Cameroons,  Gold 
Coast,  Gambia,  Senegal,  Sierra  Leone,  Lagos, 
Nigeria. 

North  Africa  :  Tripoli,  Tunis,  Algeria,  parts 
of  the  Sahara. 

Central  Africa  :  Sudan,  various  portions. 
Uganda,  Nyasaland. 

It  occurs  with  varying  frequency  in  these 
regions.  It  is  probably  more  widely  spread 
than  this  list  implies,  as  undoubtedly  many 
cases  are  seen  which  are  not  recorded. 

Isolated  cases  have  been  recorded  from 
Arabia,   India,^  Greece,  Cyprus. 

The  means  by  which  infection  is  brought 
about  are  still  uncertain  ;  we  only  know  that 
the  miracidia  (fig.  175)  enclosed  in  the  dis- 
charged eggs  do  not  hatch  if  the  eggs  remain 
in  the  urine,  but  after  cooling  perish.  As 
soon,  however,  as  the  urine  is  diluted  with 
water  the  shell  swells,  generally  bursting  length- 
ways, and  releases  the  miracidium  from  its 
investing  membrane,  so  that  it  can  swim  about 
with  the  aid  of  its  cilia.  In  its  structure  it 
differs  but  little  from  the  miracidium  of  Fasciola 
hepatica,  as,  for  instance,  in  the  lack  of  eyes  ; 
the  two  large  gland  cells  situated  on  either  side  of  the  intestinal 
sac  are  also  present  in  the  miracidia  of  Fasciola  hepatica. 

Sarcode  Globules. — This  is  a  term  applied  to  certain  globules  which  at 
times  appear  in  the  miracidium  and  are  later  ejected.     Some  authors 


V.SC. 


Fig.  1 74.  —  Schistosoma 
hamatobium^  Bilh.  :  geni- 
talia of  the  female.  V.s.^ 
ventral  sucker;  /.,  gut  caeca  ; 
V.d.^  vitelline  duct;  V.sc, 
vitellarium  ;  (?.,  ovary  ;  Oe., 
oesophagus  ;  Sh.,  shell 
gland  ;£/.,  uterus.  Magni- 
fied.    (After  Leuckart.) 


*  In  a  case  from  Madras,  recoided  by  Stephens  and  Christophers,  the  eggs  were  long  and 
spindle-shaped,  quite  unlike  the  eggs  of  Schistosoma  hcematobium. 


SCHISTOSOMA   MANSONI  277 

consider  them  as  indicative  that  the  miracidium  has  developed  into 
a  sporocyst,  but  Looss  considers  them  to  be  degeneration  products. 

The  Bilharzia  mission,  under  K.  T.  Leiper,  sent  to  Egypt  by  the 
War  Office  early  in  19 15,  reports  that  cercariae  of  bilharzia  type  were 
recognized  in  four  of  the  commonest  fresh-water  molluscs  around 
Cairo. 

With  material  obtained  from  naturally  infected  Planorbis  boissyi 
acute  bilharziosis  was  experimentally  produced  in  rats,  mice,  and 
monkeys.  Infection  takes  place  experimentally  through  the  skin  and 
also  through  the  mucous  membrane  of  the  mouth  and  oesophagus. 
The  miracidium,  after  entering  the  mollusc,  develops  into  a  sporocyst. 
This  gives  rise  not  to  rediae,   but  to  secondary  sporocysts,  which,  in 


Schistosonna  haematobium,  Bilharz,  1852. 

Male,  four  or  five  large  testes.  Gut  forks  unite  late,  so  that  the 
single  gut  stem  is  short.  Female,  ovary  in  posterior  half  of  body. 
Uterus  very  long,  voluminous,  with  many  terminal-spined  eggs,  some 
lying  in  pairs.  Vitellaria  in  posterior  fourth  of  body.  Cercariae  in 
Bullinns  contorUis  and  Bullimis  dyboivski  (syn.  :  Physa  alexandrina)  in 
Egypt. 

Schistosoma  mansoni,  Sambon,  1907. 

Male,  eight  small  testes.  Gut  forks  unite  early,  so  that  the  single 
gut  stem  is  very  long.  Females,  ovary  in  anterior  half  of  body.  Uterus 
very  short  ;  usually  only  one  lateral-spined  Qgg  at  a  time  ///  titero. 
Vitellaria  occupy  posterior  two-thirds  of  body.  Cercariae  in  Planorbis. 
boissyi  in  Egypt. 

The  above  morphological  descriptions  are  founded  on  worms  of 
each  species,  derived  from  experimentally  infected  mice  (Leiper,  R.  T., 
Brit.  Med.  Journ.y  March  18,  191 6,  p.  411). 


To  Binder  :  face  p.  276 


Syn. :  TS\  cattoi,  BlancEard,  1905. 

Male. — Eight  to  19  mm.,  but  extreme  limits  are  5  to  22*5  mm.  Con- 
sists of  a  short  fore-body,  separated  by  the  ventral  sucker  from  the 
hind-body.     The  ventral  sucker  is  stalked  and  somewhat  larger  than 


276  THE   ANIMAL   PARASITES   OF    MAN 

Besides  the  bladder,  the  ureters  and  kidneys  may  in  advanced  cases 
be  involved.  The  prostate  and  vesiciilae  seminales  are  commonly 
diseased.  Eggs  have  been  recorded  in  the  semen.  The  urethra  is 
frequently  attacked;  the  vagina  in  the  female. 

Eggs  also  occur  in  the  lymphatic  glands  of  the  gut. 

Geographical    Distribution.  —  East     Africa  : 
^-     f®^  Nil^  Valley,  Red   Sea  Coast,  Zanzibar,  Portu- 

guese East  Africa,   Delagoa  Bay,   Natal,   Port 
Elizabeth. 

South  Africa  :   Cape  Colony,  Orange  Free 
State,    Transvaal,   Mauritius,  Bourbon,  Mada- 

p-pj^c^r 


Sarcode  Globules.  —  1  nis  is  a  term  appiiea  to  certain  giODUies  wnicn  at 
times  appear  in  the  miracidium  and  are  later  ejected.     Some  authors 

*  In  a  case  from  Madras,  recoided  by  Stephens  and  Christophers,  the  eggs  were  long  and 
spindle-shaped,  quite  unlike  the  eggs  of  Schistosoma  hcematobium. 


SCHISTOSOMA   MANSON] 


277 


consider  them  as  indicative  that  the  miracidium  has  developed  into 
a  sporocyst,  but  Looss  considers  them  to  be  degeneration  products. 

The  Bilharzia  mission,  under  R.  T.  Leiper,  sent  to  Egypt  by  the 
War  Office  early  in  19 15,  reports  that  cercariae  of  bilharzia  type  were 
recognized  in  four  of  the  commonest  fresh-water  molluscs  around 
Cairo. 

With  material  obtained  from  naturally  infected  Planorbis  boissyi 
acute  bilharziosis  was  experimentally  produced  in  rats,  mice,  and 
monkeys.  Infection  takes  place  experimentally  through  the  skin  and 
also  through  the  mucous  membrane  of  the  mouth  and  oesophagus. 
The  miracidium,  after  entering  the  mollusc,  develops  into  a  sporocyst. 
This  gives  rise  not  to  redise,  but  to  secondary  sporocysts,  which,  in 
turn,  produce  cercariae.  These,  like  the  adult  worm,  differ  from  other 
distomes  in  lacking  a  muscular  pharynx. 

Schistosoma  mansoni,  Sambon,  1907. 

According  to  Manson,  Sambon  and  others,  the  eggs  with  lateral 
spines  belong  to  a  species  different  from  Schistosoma  hcBmatobium. 
Infections  with  this  species  only  are  said  to 
occur  in  the  Congo,  Southern  States  of  North 
America,  West  Indies  (Guadeloupe)  and  Brazil 
(Bahia).  The  following  characters,  according  to 
Flu,  differentiate  this  species  :  (i)  In  the  male 
the  transition  from  the  anterior  portion  of  the 
worm  to  the  lateral  fields  (the  infolded  portions 
which  form  the  gynaecophoric  canal)  is  not  a 
gradual  one  as  in  Schistosoma  hcematobium, 
but  in  this  case  the  lateral  fields  rise  suddenly, 
almost  at  right  angles  to  the  anterior  portion. 
(2)  The  ovaries  have  a  well-marked  convoluted 
course  as  in  no  other  schistosome.  (3)  The 
ootype  is  symmetrical  in  reference  to  the  long 
axis  of  the  body,  its  duct  being  lateral  on  the 
ventral  side  (Looss'  explanation  of  this  we  have 
already  given).  (4)  The  worms  live  exclusively 
in  portal  vein  and  tract.  (As  lateral-spined  eggs 
occur  also  in  the  bladder,  this  is  not  exactly  true.) 


Fig.  175.  —  Ovum  of 
Schistosoma  hamatobiuniy 
Bilh.,  with  miracidium, 
which  has  turned  its  an- 
terior end  towards  the 
posterior  end  of  the  egg. 
275/1.  (After  Looss. 


Schistosonna  japonicum,  Katsurada,  1904. 

Syn. :  S.  cattoi,  Blanchard,  1905. 

Male. — Eight  to  19  mm.,  but  extreme  limits  are  5  to  22*5  mm.  Con- 
sists of  a  short  fore-body,  separated  by  the  ventral  sucker  from  the 
hind-body.     The  ventral  sucker  is  stalked  and  somewhat  larger  than 


278 


THE   ANIMAL    PARASITES   OF   MAN 


the  oral  sucker.  Both  suckers  are  larger  than  the  corresponding  ones 
in  S.  hcematohium.  Body  usually  smooth,  but  in  the  fresh  state 
numerous  fairly  evident  spines  along  the  margin  of  the  canal.  (Eso- 
phagus :  two  bulbs.    The  junction  of  the  gut  forks  more  posterior  than 

in  S\  hcematohhmij  the  median  united  gut 
stem  occupying  a  quarter  to  one-fifth  to 
one-sixth  of  the  body  length.  An  excre- 
tory canal  runs  along  each  side  of  the 
body,  opening  into  the  dorsal  excretory 
pore.  Testes  irregularly  elliptical,  six  to 
eight  in  number,  in  the  anterior  part  of 
hind-body.  The  vasa  efferentia  unite 
into  a  common  vas  deferens  which  opens 
directly  behind  the  ventral  sucker.  The 
seminal  vesicle  lies  just  behind  this. 

Female. — Up  to  26  mm.,  generally 
thinner  than  the  male.  Surface  smooth. 
Suckers  armed  with  fine  spines.  Ventral 
sucker  larger  than  oral.  Body  thicker 
behind  the  region  of  the  ovary.  The 
gut  forks  unite  immediately  behind  the 
ovary.  The  united  gut  much  thicker 
than  in  S.  liceinatobiujii.  Ovary  elliptical, 
almost  in  the  mid-body,  its  hinder  por- 
tion dilated.  The  oviduct  arises  from 
its  posterior  end  and  then  runs  sinuously 
forward,  where  it  is  joined  by  the  vitel- 
larian  duct ;  the  vitellarium  well  deve- 
loped, extending  from  behind  the  ovary 
almost  but  not  quite  to  the  posterior 
end  as  in  S.  hctmatobitun.  Shell  gland 
ducts  enter  at  the  junction  point  of 
oviduct  and  vitelline  duct.  The  canal 
here  forms  an  ootype  and  then  proceeds 
as  the  uterus  to  open  directly  behind 
the  ventral  sucker.  The  uterus  occupies 
almost  half  the  hind-body.  In  S\  hcema- 
tobiiim  this  is  not  so.  The  uterine  canal 
is  cleft-like,  i.e.,  its  dorso-ventral  diameter 
is  much  greater  than  its  lateral  diameter. 
The  number  of  eggs  varies  from  about 
50  to  300  from  observations  made  in  various  hosts. 

Eggs. — In  iitero  assume  various  shapes,  as  they  are  soft ;  the  lumen 
of  the  uterus  is  narrow.     Outside  they  are  oval,  faint  yellow,  double 


Fig.  176. — Schistosoma  japoni- 
cum  :  anterior  end  with  testes  ; 
posterior  end  with  point  of  union 
of  caeca.  Length  of  worm  about 
10  mm.     (After  Katsurada.) 


SCHISTOSOMA  JAPONICUM 


279 


contoured.     In  faeces  the  eggs  measure  83*5 /^  by  62*5/1  (man) ;  85 /x 
by  61-5  ^  (cattle)  ;  98*2  fju  by  73*8  fi  (dog).     The   eggs  have   either 
small    lateral    spines    or    thickenings,    and    Looss    at    the    opposite 
side     has    described    cap-like    thickenings. 
The  eggs  in  the  tissues  undergo  various  de- 
formities, and  may  contain  a  miracidium,  as 
also  the  eggs  in  faeces  do  ;    or  the  contents 
may  consist  of  granular  matter  or  amorphous 
masses  or  they  may  be  calcified.     Lympho- 
cytes and  giant  cells  may  also   invade  the 
eggs. 

Mode  of  Infect  ion. ^The  miracidia  hatch 
in  water  in  as  little  as  fifteen  minutes,  but 

the  majority  in  one  to  three  hours.  They  will  live  in  water  for  about 
twenty-four  hours.  In  water  they  undergo  a  transformation  into 
*'  larvae/'    which    then    penetrate   the    skin,    as    has   been  shown  by 


Fig.  177. — Schistosoma  japo- 
nicum,  male  and  female  in  co- 
pula.  X  60.  (After  Katsurada.) 


Fig.    178. — Schistosoma  japonicum  :  eggs  from  human  liver,  showing  "spines"  and 
"hoods"  at  opposite  pole.     (After  Looss.) 


Japanese  writers  to  hold  good  for  man,  cattle,  dog  and  cat.  The 
penetration  of  the  skin  is  attended  with  an  eruption  on  the  legs, 
"■  Kabure."  The  exact  route  by  which  the  worms  reach  the  portal  vein 
is  uncertain.  Infection  in  Japan  takes  place  from  spring  to  autumn, 
especially  May  to  July,  when  the  soil  is  contaminated  with  manure 
of  cattle  infected  with  S.  japonicum.  They  also  appear  to  develop 
in    molluscs.     Leiper   and  Atkinson    found    cercariae  (in   sporocysts) 


28o 


THE   ANIMAL   PARASITES   OF    MAN 


in  the  liver  of  a  mollusc,  Katayama  nosophora.  They  infected  mice 
by  immersing  them  in  water  containing  liver  emulsion  and  so  free 
cercariae,  thus  confirming  the  similar  results  of  Miyairi  and  Suzuki. 

Habitat. — The  worm  occurs  in  Japan,  China,  and  the  Philippines. 
The  normal  host  is  man  and  mammals.  Cattle,  dog  and  cat  are 
often  found  naturally  infected.  Mice  can  also  be  experimentally 
infected.     Their  seat  of  election  is  the  portal  vein  and  its  branches, 

especially  the  mesenteric  veins.  They  either 
swim  free  in  the  blood  or  remain  fixed  by 
their  suckers  to  the  intima  of  the  vessels. 
They  have  also  been  found  in  the  vena  cava 
and  right  heart  of  a  cat,  but  not  so  far  in  the 
vesical  plexus. 

Eggs  are  found  in  the  submucosa  and 
mucosa  of  the  gut,  especially  the  colon,  and 
at  times  in  the  serosa  and  subserosa  of  the 
small  intestine,  where  they  give  rise  to  new 
growths.  Occasionally  eggs  are  found  in 
the  brain.  The  life  of  tlie  worms  is  at  least 
two  years. 

Pathogenic  Effects.  —  An?emi3.  through  loss 

of  blood  due   to    worms  ;    enlarged  spleen, 

toxic  in  origin  (?) ;  phlebitis,  thrombosis,  due 

to  portal  stasis  ;  the  eggs,  however,  cause  the 

They  are   carried  by   the   circulation    to   various 


Fig.  179. — Schistosoma  Ja- 
ponicum  :  from  dog.  Uterine 
^gg*  "^  c.  800.  (After  Kat- 
surada.) 


greatest  mischief. 


7/ 


Fig.    v%0.~ Schistosoma   japonictim  :  from 
dog.      X  c.  800.     (After  Katsurada.) 


Fig.  181. — Schistosoma  japonicum  : 
from  dog.  Egg  from  faeces,  x  c.  Scx). 
(After  Katsurada.) 


SCHISTOSOMA   JAPONICUM  281 

organs  where  they  produce  inflammation,  granulation  tissue,  and  later 
connective  tissue. 

Liver. — The  eggs  reaching  this  organ  give  rise  to  granulomata  and 
hence  enlarged  liver,  and  later,  when  connective  tissue  is  formed,  to 
contraction.  The  surface  is  rough  and  irregularly  granular,  ''parasitic 
embolic  cirrhosis  "  of  Yamagiwa. 


^^ 


rf% 


ms€>  Q 


0  (■    ■  o-}V>xr^^ 


■n 


^ 


^>-\>-;    rPS^c;,..,        -      :     ^c^ 


O' 


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o 


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'■■  .J 

Fig.    \%2.— Schistosoma  japonicum  :  section    through  the  gut  of  a  Chinaman  showing  eggs. 

X  58.     (After  Catto.) 

Gut. — The  eggs  in  the  mucosa  and  submucosa  cause  catarrh  and 
destruction  of  tissue  or  new  growth.  In  the  small  intestine  the  eggs 
are  mainly  in  the  serosa  and  subserosa,  where  they  give  rise  to  poly- 
poid or  branched  growths. 


282 


THE   ANIMAL   PARASITES   OF   MAN 


S/)/^^n.— Enlarged,  at  first  due  to  toxin  (?)  and  later  due  to  portal 
stasis.     Eggs  in  the  spleen  are  uncommon. 

Ascites  also  arises  from  the  portal  stasis,  and  is  generally  present 
in  advanced  cases. 

Eggs  may  be  found  in  many  other  situations  :  glands  (numerous), 
mesentery,  stomach,  pancreas,  kidney,  etc.    The  bladder  remains  free. 


Fig.  183. — Schistosoma  japonicuni  :  liver  showing  eggs  in  the  intra-  and   interlobular 
connective  tissue,      x    c.  80.     (After  Katsurada.) 


Class  III.     CESTODA,  Rud.,  1808. 


Tapeworms  have  been  known  from  ancient  times — at  all  events,  the  large 
species  inhabiting  the  intestines  of  man — and  there  has  never  been  a  doubt  as 
to  their  animal  nature.  The  large  cysticerci  of  the  domestic  animals  (occasionally 
of  man  also)  have  been  known  for  an  equally  long  period,  but  they  were  generally 
regarded  as  growths,  or  "hydatids,"  until  almost  simultaneously  Redi  in  Italy,  and 
Hartmann  and  Wepfer  in  Germany,  concluded  from  their  movements  and  organi- 
zation that  they  were  of  animal  nature.  From  that  time  the  cysticerci  have  been 
included  amongst  the  other  intestinal  worms,  and  Zeder  (1800)  established  a  special 
class  {Cystici,  Rud.,  1808)  for  the  bladder  worms.  Things  remained  in  this  condition 
until  the  middle  of  the  last  century,  when  Kiichenmeister,  by  means  of  successful 
feeding  experiments,  demonstrated  that  the  cysticerci  were  definite  stages  of 
development  of  certain  tapeworms.  Before  Kiichenmeister,  E.  Blanchard,  van 
Beneden,  and  v.  Siebold  had  held  the  same  opinion  in  regard  to  other  asexual 
Cestodes. 


CESTODA  283 

Since  the  most  remote  period  another  question  has  again  and  again  occupied 
the  attention  of  naturahsts,  the  question  of  the  morphological  nature — that  of  the 
INDIVIDUALITY  OF  THE  TAPEWORM.  The  ancients,  who  were  well  acquainted 
with  the  proglottids  {Vermes  aecurbitani)  that  are  frequently  evacuated,  were 
of  the  opinion  that  the  tapeworm  originated  through  the  union  of  these  separate 
proglottids,  and  this  view  was  maintained  until  the  end  of  the  seventeenth 
century.  In  1683  Tyson  discovered  the  head  with  the  double  circlet  of  hooks  in 
a  large  tapeworm  of  the  dog  ;  Redi  (1684)  was  also  acquainted  with  the  head  and 
the  suckers  of  several  Tsenice.  Andry  (1700)  found  the  head  of  Tcenia  saginata^  and 
Bonnet  (1777)  and  Gleichen-Rusworm  (1779)  found  the  head  of  Diboihriocephalus 
latus.  Consequently  most  authors,  on  the  ground  of  this  discovery,  considered  the 
tapeworm  as  a  single  animal,  that  maintains  its  hold  in  the  intestine  by  means  of  the 
head,  and  likewise  feeds  itself  through  it.  The  fact  was  recognized  that  there  were 
longitudinal  canals  running  through  the  entire  length  of  the  worm,  and  it  was  thought 
that  these  originated  in  the  suckers,  and  that  the  entire  apparatus  was  an  intestine. 
As,  moreover,  the  segments  form  at  the  neck,  and  are  cast  off  from  the  opposite 
extremity,  the  tapeworm  was  also  compared  with  the  polyps,  which  were  formerly 
regarded  as  independent  beings. 

Steenstrup,  in  his  celebrated  work  on  the  alternation  of  generations  (1841),  was 
the  first  to  give  another  explanation.  This  has  been  elaborated  still  further  by 
van  Beneden,  v.  Siebold  and  Leuckart,  and  until  a  few  years  ago  all  authorities 
adopted  his  views.  According  to  this  view,  the  tapeworm  is  composed  of  numerous 
individuals,  something  like  a  polyp  colony,  and,  in  addition  to  the  proglottids — the 
sexual  individuals  which  are  usually  present  in  large  numbers — there  is  ONE 
individual  of  different  structure,  the  scolex^  which  not  only  fastens  the  entire  colony 
to  the  intestine,  but  actually  produces  this  colony  from  itself,  and  therefore  is 
present  earlier  than  the  proglottids.  The  scolex  is  a  "nurse,"  which,  though  itself 
produced  by  sexual  means,  increases  asexually  like  a  Scyphistoma  polyp  ;  the 
tapeworm  chain  has  therefore  been  termed  a  strobila.  Consequently  the  develop- 
ment of  the  tapeworms  was  explained  by  an  alternation  of  generations.  In  support 
of  this  opinion  it  was  demonstrated  not  only  that  the  adult  sexual  creatures,  the 
proglottids,  can  separate  from  the  colony  and  live  independently  for  a  time,  but 
that  in  certain  Taeniae,  and  especially  in  many  Cestodes  of  the  shark,  the  proglottids 
detach  themselves  long  before  they  have  attained  their  ultimate  size,  and  thus 
separated  continue  to  develop,  grow  and  finally  multiply  ;  the  scolex  also  exhibits  a 
certain  independence  in  so  far  as,  though  not,  as  a  rule,  capable  of  a  free  life,  yet 
it  in  some  cases  lives  as  a  free  being,  partly  on  the  surface  of  the  body  of  marine 
fishes  and  partly  in  the  sea.  With  the  more  intimate  knowledge  of  the  develop- 
ment of  the  cysticerci,  the  independent  nature  of  the  scolex  was  recognized.  It  is 
formed  by  a  budding  of  the  bladder  that  has  developed  from  the  oncosphere,  in 
some  cases  (Coenurus)  in  large  numbers,  in  other  cases  (Echinococcus)  only  after 
the  parent  cyst  has  developed  several  daughter  cysts.  Released  from  its  mother 
cyst  and  placed  in  suitable  conditions,  it  goes  on  living,  and  gives  rise  at  its 
posterior  end  by  budding  to  the  strobila,  the  proglottids  of  which  eventually  become 
sexual  individuals. 

In  order  to  make  this  clearer  we  will  briefly  summarize  what  takes  place  in 
the  jelly-fishes. 

V>ymetamorphosis  is  meant  a  developmental  change  in  the  sa7ne  indivi'dual,  while 
alternation  of  generations,  or  metagenesis,  implies  a  stage  in  which  reproduction  of 
individuals  takes  place  by  a  process  of  budding  or  fission.  This  asexual  reproduc- 
tive stage  alternates  with  the  sexual  mode  of  reproduction.  Thus  in  the  development 
of  the  Scyphozoa  (jelly-fishes)  we  have  : — 

(r)  The  fertilized  ^g%  cell  divides  regularly  and  forms  a  morula. 


284  THE   ANIMAL   PARASITES   OF   MAN 

(2)  By  accumulation  of  fluid  in  the  interior  this  becomes  a  closed  sac  with  a  wall 
formed  of  a  single  layer  of  cells,  forming  the  blastosphere  or  blastula. 

(3)  One  end  of  the  sac  is  invaginated,  forming  a  gastrula. 

(4)  The  gastrula  pore  or  mouth  closes,  forming  again  a  sac,  the  walls  of  which 
have  two  layers,  forming  2iplanula. 

(5)  This  becomes  fixed  to  a  rock,  an  invagination  forms  at  one  end,  a  depression 
— the  stomodaeum — communicating  with  the  enteric  cavity.  Tentacles  grow  out 
and  we  have  a  Scyphozobn  polype^  Scyphistoma  or  Scyphula.  It  is  to  this  stage  that 
Steenstrup  gave  the  name  "  nurse  "  ("  wet-nurse  ")»  because  it  nourished  or  produced 
asexually  the  succeeding  forms. 

(6)  Asexual  reproduction  by  transverse  fission  occurs  in  this,  forming  a  pile  of 
saucer-  or  pine-cone-like  animals  which  before  this  time  had  been  considered  to 
be  a  distinct  animal,  which  was  called  sirobila  from  its  resemblance  to  a  pine- 
cone.     This  is  the  alternate  generation. 

(7)  The  individuals  of  the  strobila  become  free  and  are  called  Epkyrulcs. 

(8)  These  develop  finally  into  adult  sexual  jelly-fish,  Scyphozoa,  so  that  com- 
paring a  tapeworm  with  this  we  have  {a)  t^g,  {b)  scolex  (=  Scyphula  or  "  nurse  "), 
{c)  asexual  reproduction  of  the  tapeworm  chain  (=  strobila),  id)  development  of  the 
individuals  of  the  chain  (proglottids)  into  sexual  adults. 

Van  Beneden's  terminology  for  these  stages  is  the  following :  Ciliated  embryo 
=  protoscolex  ;  scyphistoma  —  deutoscolex  (or  scolex)  ;  free  Ephyrula  =  proglottis. 
According  to  this  view,  as  is  the  case  in  many  endoparasitic  Trematodes,  asexual 
reproduction  by  budding  occurs  at  two  stages  of  the  whole  cycle  of  development, 
viz.  (i)  in  the  formation  of  the  scolex  by  budding  from  the  bladder  ("nurse"), 
(2)  in  the  formation  of  the  strobila  by  budding  from  the  scolex  ("nurse"). 

But  in  cysticercal  larval  forms  it  appears  that  the  scolex  does  not  arise  in  this 
way  but  is  simply  a  part  of  the  proscolex  (hexacanth  embryo),  becoming  invaginated 
into  it  for  protection,  so  that  there  is  no  asexual  gemmation  here.  It  has  been 
questioned  also  whether  the  strobila  also  arises  by  gemmation.  If  it  does,  the 
tapeworm  is  a  colony  of  zooids  produced  by  budding  from  the  asexual  scolex  ;  if  it 
is  not  produced  in  this  way,  then  the  tapeworm  is  to  be  regarded  as  an  individual 
in  which  growth  is  accompanied  by  segmentation.  Against  the  "  colony "  view  are 
the  facts  that  the  muscular,  nervous,  and  excretory  systems  are  continuous  through- 
out the  worm,  and  that  some  tapeworms,  such  as  Ligula^  are  unsegmented. 

Finally,  if  the  tapeworm  is  an  individual  the  question  arises  which  is  the  head 
end.  As  new  segments  are  formed  at  the  neck,  and  as  this  point  in  annelids  is 
the  antepenultimate  segment,  the  scolex  must  be  the  last  or  posterior  segment.  The 
caudal  vesicle  or  bladder  of  larval  forms  is  consequently  anterior.  According  to 
this  view,  in  tapeworms  as  among  many  endoparasitic  flukes,  an  asexual  multiplica- 
tion occurs  at  two  points  of  the  whole  cycle  of  development,  which  is  as  follows : 
(i)  ^%%^  (2)  oncosphere  or  hexacanth  embryo,  (3)  bladder  (cysticercus  or  hydatid), 
(4)  (after  digestion  of  the  bladder)  by  budding,  the  scolex,  (5)  by  budding  from  the 
scolex  the  sexual  proglottids,  (6)  the  %%%  ;  (4)  and  (5)  being  the  two  asexual  stages. 

Anatomy  of  the  Cestoda. 

If  we  except  the  tapeworms  with  only  one  proglottis,  the 
Cestoidea  Monozoa,  Lang  =  Cestodariay  MonticeUi,  we  can  always 
distinguish  in  the  Cestodes,  in  the  narrower  sense,  one  scolex  or  head 
and  a  large  or  small  number  of  segments  (proglottids).  The  scolex 
serves  the  entire  tapeworm  for  fastening  it  to  the  internal  surface  of  the 


ANATOMY   OF    THE    CESTODA  285 

intestinal  wall,  and  therefore  carries  at  its  end.  various  organs  which 
assist  in  this  function,  and  which  are  as  follows  :  (i)  Suctorial 
ORGANS,  i.e.,  the  four  suckers  (acetabula),  which  are  placed  crosswise 
at  the  circumference  of  the  thickened  end  of  the  scolex  ;  further,  the 
double  or  quadruple  groove-like  suckers  (bothridia),  which  are 
diversely  shaped  in  the  various  genera  and  families/  (2)  Fixation 
ORGANS  (hooklets)^  that  likewise  occur  in  varying  numbers  and 
different  positions;  they  may  be  in  the  suckers,  or  outside  them  on  the 
apex  of  the  scolex  ;  for  instance,  in  many  of  the  Tceniidce  they  appear 
in  a  circle  around  a  single  protractile  organ,  the  rostellum,  or  the  latter 
may   be   rudimentary,    and   is   then    replaced   by  a   terminal  sucker. 

(3)  Proboscis.  One  family  of  the  Cestodes,  the  RhynchobothriidcB, 
carries  four  proboscides,  moved  by  their  own  muscular  apparatus, 
on    the   scolex,   and    they   are    beset  with   the   most   diverse    hooks. 

(4)  Tentacle-like  formations  are  only  known  in  one  genus 
(Polypocephalus). 

The  thickened  part  of  the  scolex  that  carries  the  suckers  is  usually 
called  the  head  ;  the  following  flat  (unsegmented)  part  connecting  it 
with  the  proglottids  is  called  the  neck,  and  is  sometimes  quite  small. 
In  a  few  cases  the  entire  scolex  (or  head)  disappears,  and  its  function  is 
then  undertaken  by  the  contiguous  portion  of  the  chain  of  proglottids, 
which  is  transformed  into  a  variously  shaped  PSEUDO-SCOLEX. 

The  proglottids  are  joined  to  the  scolex  in  a  longitudinal  row, 
and  are  arranged  according  to  age  in  such  a  manner  that  the  oldest 
proglottis  is  farthest  from  the  scolex,  and  the  youngest  nearest  to  it. 

The  number  of  segments  varies,  according  to  the  species,  from 
only  a  few  to  several  thousands ;  they  are  either  quadrangular  or 
rectangular ;  in  the  latter  case  their  longitudinal  axis  falls  either 
longitudinal  or  transverse  to  that  of  the  entire  chain,  according  as 
the  segments  are  longer  than  broad  or  broader  than  long.  When  the 
number  of  segments  is  very  large,  the  youngest  ones  are,  as  a  rule, 
transversely  oblong,  the  middle  ones  are  squarish,  and  the  mature 
ones  longitudinally  oblong.  The  posterior  border  of  the  segments, 
as  a  rule,  carries  a  longitudinal  groove  for  the  reception  of  the  shorter 
anterior  border  of  the  following  proglottis.  The  two  lateral  borders 
of  the  segment  are  rectilinear,  but  converge  more  or  less  towards  the 
front,  or  they  are  bent  outwards.  In  most  of  the  Cestodes  the 
segments,    just    as    the    neck,    are    very    fiat  ;    in    rare    cases    their 

1  They  may  remain  simple,  and  are  then  not  separated  from  the  remaining  muscles 
of  the  scolex  ;  or  they  project  as  roundish  or  elongated  structures  over  the  scolex,  hollow  on 
their  free  surface,  and  often  divided  into  numerous  areas  by  muscular  transverse  ribs.  They 
may  also  carry  accessory  suckers  on  their  surface. 

2  The  various  parts  of  a  booklet  are  thus  named  from  the  point  backwards :  (i)  blade  or 
prong,  (2)  guard  or  ventral  or  posterior  root,  (3)  handle  or  dorsal  or  anterior  root. 


286  THE   ANIMAL    PARASITES   OF   MAN 

transverse  diameter  is  equal  to  their  dorso-ventral  diameter.  As  a 
rule  the  segments,  singly  or  several  united  together,  detach  them- 
selves from  the  posterior  end,  in  many  cases  only  after  complete 
maturity  is  attained,  and  in  others  much  earlier ;  they  then  continue 
to  live  near  their  parent  colony,  to  still  call  it  by  that  name,  in  the 
same  intestine  and  continue  their  development.  Even  when  evacuated 
from  the  intestine  the  proglottids  under  favourable  circumstances 
can  continue  to  live  and  creep  about,  until  sooner  or  later  they 
perish. 

The  first  proglottis  formed,  and  which  in  a  complete  tapeworm 
[i.e.,  sexually  complete]  is  the  most  posterior,  is  as  a  rule  smaller  and 
of  different  shape,  it  also  frequently  remains  sterile,  as  likewise  happens 
in  the  next  (younger)  segments  in  a  few  species ;  otherwise,  however, 
sooner  or  later  the  generative  organs  develop  in  all  the  segments, 
mostly  singly,  sometimes  in  pairs ;  in  the  latter  case  they  may  be  quite 
distinct  from  each  other  or  possess  some  parts  in  common.  The 
term  *'  mature "  is  used  for  a  proglottid  that  has  the  sexual  organs 
fully  developed,  while  '^gravid"  is  used  for  one  containing  eggs. 
Most  of  the  species  combine  male  and  female  genitalia  in  the  same 
segment,  only  a  few  are  sexually  distinct  (Dioecocestus).  In  the 
hermaphrodite  species  one  male  and  one  female  sexual  orifice  are 
always  present,  and,  in  addition,  there  may  be  a  second  female  orifice, 
the  uterine  opening ;  as  a  rule,  however,  this  is  lacking,  and  in  one 
sub-family,  the  A  cole ince,  to  which  also  the  genus  Dioecocestus  belongs, 
the  other  sexual  orifice,  the  opening  of  the  vagina,  is  also  absent. 
The  position  of  these  orifices  varies  ;  the  cirrus  and  vagina  usually 
open  into  a  common  atrium  on  one  lateral  border  or  on  a  surface 
of  the  segments ;  the  orifice  of  the  uterus  may  be  on  the  same 
surface  or  on  the  opposite  one. 

The  surface  on  which  the  uterus  opens  is  termed  the  ventral 
SURFACE;  if  this  orifice  is  absent,  one  must  depend  on  the  ovary, 
which  almost  always  approaches  one  of  the  two  surfaces ;  this  surface 
is  then  called  the  ventral. 

The  length  of  the  Cestodes — independently  of  their  age — depends 
on  the  number  and  size  of  the  segments,  as  well  as  on  their 
contraction  ;  the  smallest  species  {Davainea  proglottina)  is  0*5  to 
vo  mm.  in  length;  the  largest  may  attain  a  length  of  10  m.,  and 
even  more. 

The  entire  superficial  surface  of  the  tapeworms  is  covered  with 
a  fairly  resistant  and  elastic  layer,  which  exhibits  several  indistinctly 
limited  layers  and  which  is  usually  called  a  cuticle,  which  also  covers 
the  suckers,  and  is  reflected  inwardly  at  the  sexual  orifices.  In  some 
species  fine  hairs  appear,  either  on  the  entire  body  or  only  in  the 
region  of  the  neck,  on  the  external  surface.     In  the  cuticle  there  can  be 


ANATOMY  OF  THE  CESTODA 


287 


A.m. 


recognized,  besides  the  pores,  which  no  doubt  are  concerned  with 
nutrition,  spaces  in  which  He  the  ends  of  sensory  cells.  Close  under 
the  cuticle  lies  the  external  layer  of  the  parenchyma  (basal  membrane), 
and  below  this  the  circular  and  longitudinal  muscles  forming  the 
dermo-muscular  coat.  The  matrix  cells  of  the  cuticle  occur  as  in 
the  Trematodes,  only  on  the 
inner  side  of  the  peripheral 
muscles  in  the  external  zone 
of  the  parenchyma  ;  they  are 
fusiform  cells,  forming  one 
or  two  layers,  but  are  not 
arranged  in  the  manner  of 
epithelial  cells  (fig.  184,  Sec). 
They  have  fine  branching 
processes  which  run  between 
the  dermal  muscles,  pass 
through  the  basal  membrane 
and  penetrate  the  internal  sur- 
face of  the  cuticle  with  small 
pistil-like  enlargements,  ex- 
panding on  the  internal 
surface  of  the  cuticle  into 
a  thin  plasma  layer. 

In  addition  to  the  above 
mentioned,  there  are  other 
cuticular  formations  occur- 
ring on  the  cuticle  of  some 
Cestodes,  such  as  immobile 
hairs  and  variously  formed 
hooks,  such  as  are  seen  prin- 
cipally on  the  scolex.  Their 
development  is  only  roughly 
known  in  a  few  species  ;  they 
are  usually  already  present  in 
the  larval  stage,  and  of  the 
same  arrangement  and  shape 
as  in  the  fully  developed 
tapeworms  ;  a  matter  of 
importance,  because  by  these 


Ex.c. 


F.v.s. 


Fig.  184. —  Schematic  representation  of  a  small 
part  of  a  transverse  section  o{  Ligula  sp.  Bs.,  basal 
membrane  ;  Cn.,  cuticle  ;  at  its  base  are  the  end- 
plates  of  the  subcuticular  (epithelial)  cells;  in  the 
centre  a  cuticular  sense  organ,  O.s.  ;  F.v.s.,  vitelline 
follicle  ;  Exc,  excretory  vessel  ;  C,  calcareous  cor- 
puscle ;  L.m.,  longitudinal  muscles  ;  M.c.  myoblast ; 
P.m.,  parenchymatous  or  dorso-ventral  muscles  ; 
/*/.,  plexus  of  nerve  fibres  ;  A.m.,  circular  muscles; 
Sc.c.y  subcuticular  or  matrix  cell;  T.c,  terminal 
flame  cell.     500/1.     (After  Blochmann.) 


structures  larvae  can  be  recog- 
nized as  being  those  of  a  certain  species  of  tapeworm. 

The  CUTICULAR  GLANDS  in  Cestodes  are  scarce. 

The  PARENCHYMA  forms  the  chief  tissue  of  the  entire  body,  and  in 
all  essentials  its  structure  is  similar  to  that  of  the  Trematodes. 


288 


THE   ANIMAL   PARASITES   OF   MAN 


The  same  doubt  exists  here  also  as  to  the  nature  of  the  parenchyma. 
Recent  authors  consider  that  it  consists  of  highly  branched  cells,  the 
processes  of  which  ramify  in  all  directions.  These  cells  lie  in  a  non- 
cellular  matrix  containing  fluid  vacuoles.  This  matrix  spreads  in 
between  and  so  breaks  the  continuity  of  the  epidermal  cells. 

In  the  parenchyma  of  almost  all  the  Cestodes  there  are  found  in 
adult  specimens,  as  well  as  in  larvae,  light-refracting  concentrically 
striated  structures,  of  a  spherical  or  broad  elliptical  shape,  which,  on 
account  of  their  containing  carbonate  of  lime,  are  termed  calcareous 
CORPUSCLES  (fig.  184,  C).  Their  size,  between  3  ^  and  30  ^,  varies 
according  to  the  species  ;  their  frequency  and  distribution  in  the  paren- 
chyma  also  varies,  but  they  are  chiefly  found  in  the  cortical  layer. 


L.m. 


Sec. 


M.f. 


L.m. 


Sm.f. 


Fig.  185. — Half  of  a  transverse  section  through  a  proglottis  of  Tcenia  crassicollis. 
Cu.,  cuticle;  Ex.v.,  external  excretory  vessel,  to  the  right  of  which  there  is  the  smaller 
internal  one ;  T.,  testicular  vesicles  ;  L.m.  longitudinal  muscles  (outer  and  inner) ;  M./.,  lateral 
nerve  with  the  two  accessory  nerves;  Sec,  subcuticular  matrix  cells;  Sm.f.,  submedian 
nerve;  Tr.m.,  transverse  muscles;  67.,  the  uterus,  and  the  middle  of  the  entire  transverse 
section.     44/1. 

They  are  the  product  of  certain  parenchymatous  cells,  in  the  interior 
of  which  they  lie  like  a  fat  globule  in  a  fat  cell,  but  according  to 
others  they  are  intercellular  in  origin. 

The  MUSCULAR  SYSTEM  of  the  proglottids  is  composed  of— (i)  the 
subcuticular  muscles  (figs.  184  and  185),  as  a  rule  consisting  of  a 
single  layer  of  annular  muscles  ;  (2)  longitudinal  muscles;  (3)  dorso- 
ventral  fibres  extending  singly  from  one  surface  to  the  other,  and 
at  both  ends  expanding  in  a  brush-like  manner,  and  inserted  into 
the  basal  membrane,  consisting  of  an  outer,  more  numerous,  and  an 
inner,  less  numerous  but  more  powerful  layer  (the  number  of  bundles 
in  this  layer  being  in  certain  cases  of  specific  importance)  ;  (4)  trans- 
verse fibres,  the  elements  of  which  penetrate  to  the  borders  of  the  seg- 
ments, thus  passing  through   the  longitudinal  muscles   and  reaching 


ANATOMY    OF  THE   CESTODA 


289 


.--C 


the  cuticle.      In    the  region  of  the  septa  the  transverse  and  dorso- 
ventral  muscles  form  a  kind  of  plate. 

The  mass  of  parenchyma  bounded  by  the  transverse  muscles  is 
termed  the  medullary  layer,  while  the  mass  lying  outside  them 
is  termed  the  CORTICAL  layer. 

It  was  known  long  ago  that  the  myoblasts  adhere  to  the  dorso- 
ventral  fibres  as  thickenings,  but  it  is  only  recently  that  large  star- 
shaped  cells  (fig.  184),  separated  from  but  connected  with  them  by 
processes,  have  been  recognized  as 
the  myoblasts  of  other  fibres  (Bloch- 
mann,  Zernecke). 

Within  the  scolex  the  direction 
and  course  of  the  muscular  layers 
change. 

The  suckers  are  parts  of  the 
musculature,  locally  transformed, 
with  a  powerful  development  of 
the  dorso -ventral  muscles,  now 
become  radial  fibres. 

The  rostellum  of  the  armed 
Taeniae,  like  the  proboscis  of  the 
Rhynchobothfiidce,  also  belongs  to 
the  same  category  of  organs. 

In  the  simplest  form,  the  ros- 
tellum, or  top  of  the  head  (as 
in  Dipylidiuin  caninum),  appears 
as  a  hollow  oval  sac,  the  anterior 
part  of  which,  projecting  beyond 
the  upper  surface  of  the  head,  carries  several  rows  of  hooks  (fig.  186). 
The  entire  internal  space  of  the  sac  is  occupied  by  an  elastic,  slightly 
fibrous  mass,  while  the  anterior  half  of  the  surface  of  the  rostellum  is 
covered  by  longitudinal  fibres  and  the  posterior  half  by  circular  fibres. 
On  contraction  of  the  latter  the  entire  mass  is  protruded  through  the 
apical  aperture,  the  surface  of  the  rostellum  becomes  more  arched,  and 
the  position  of  the  hooks  is,  in  consequence,  altered.  The  rostellum 
of  the  large-hooked  Ta^niidcv,  which  inhabit  the  intestine  of  man  and 
beasts  of  prey,  is  of  a  far  more  complicated  structure,  for,  in  addition 
to  the  somewhat  lens-shaped  rostellum  carrying  the  hooks  on  its  outer 
surface,  there  are  secondary  muscles  grouped  in  a  cup-like  manner 
(fig.  187).  Every  change  in  the  curvature  of  the  surface  of  the  rostellum 
induces  an  alteration  in  the  position  of  the  hooks.  In  the  hookless 
Tctniidct  the  muscular  system  of  the  rostellum  is  altered  in  a  very 
different  manner  ;  in  a  few  forms  a  typical  sucker  appears  in  its  place. 
The  NERVOUS  system  commences  in  the  scolex  and  runs  through 

19 


Fig.  186. — Dipylidium  caninum  :  from 
the  cat.  In  the  upper  figure  the  rostellum  is 
retracted,  in  the  lower  protruded,  a,  sucker  ; 
<5,  hooks  of  rostellum  ;  B,  enlarged  hook  ; 
c,  apical  aperture  on  scolex  ;  d,  longitu- 
dinal muscles;  (?,  circular  muscles.  (After 
Benham. ) 


290 


THE   ANIMAL   PARASITES   OF   MAN 


the  neck  and  the  entire  series  of  proglottids.  Within  the  proglottids 
it  consists  of  a  number  of  longitudinal  nerve  fibres  of  which  those  at 
each  lateral  border  are  usually  the  largest.     In  the  Taeniae  the  lateral 

nerves   are   accompanied  both 
dorsally    and    ventrally    by    a 
thinner  nerve  (accessory  nerve) 
(fig.    185)  ;    on    each    surface, 
moreover,   between  the  lateral 
nerve   and   the  median  plane, 
there     are     two    somewhat 
stronger  bundles  (sub-median), 
so  that  there  is  a  total  of  ten 
longitudinal      nerve     bundles. 
They  lie  externally  to  the  trans- 
verse   muscle  plates,    and   the 
lateral  and  accessory   bundles 
lie   externally  to  the  principal 
excretory  vessels,  and  are  every- 
where connected  by  numerous 
anastomoses      and     secondary 
anastomoses ;  one  typical  ring 
commissure    is    usually   found 
at  the  posterior  border  of  the 
segments.     ^In     the     Boihrio- 
cephalidcv   the    distribution    of 
the  nerve  bundles  is  different 
(for   instance,    two   lie    in    the 
medullary   layer),  or  they  are 
split  up  into  a  larger  number 
of  branches.     In  the  scolex  the  nerve  bundles  are  connected  in  a  very 
remarkable    manner    by   commissures   with    that  which    is   generally 
termed  the  central  part  of  the  entire  nervous  system.     There  occurs 
normally  a  commissure  between  the  two  lateral  nerves  ;  at  the  same 
level,  the  dorsal  and  ventral  median  nerves  are  also  connected  at  each 
surface  as  well  with  each  other^^as  with  the  lateral  nerves,  so  that  a 
hexagonal  or  octagonal  figure  is  formed.     The  so-called  apical  nerves 
pass  from  this  commissural  system  anteriorly,  embrace  the  secondary 
muscular  system  of  the  rostellum  semicircularly,  and  form  an  annular 
commissure  (rostellar  ring)  at  the  inner  part  of  the  rostellum. 

The  peripheral  nerves  arise  from  the  nerve  bundles  as  well  as 
from  the  commissures  situated  in  the  scolex  ;  some  go  direct  to  the 
muscles,  while  others  form  a  close  plexus  of  nerves  external  to  the 
inner  longitudinal  muscles,  which  plexus  likewise  sends  out  fibres  to 
the    muscles,    but    principally    to  [{numerous   fusiform   sense   organs 


Fig.  187. — Longitudinal  section  of  the  head 
and  neck  of  Tienia  crassicollis,  showing  the 
lens-shaped  muscular  rostellum,  with  two  hooks 
lying  in  the  concentric  cup-like  mass  of  muscles. 
L.f/i.,  longitudinal  muscles  of  the  neck  ;  Lf.,  left 
lateral  nerve  ;  C,  ganglion  ;  S.c,  subcuticular 
layer ;  W^,  external,  W^,  internal  excretory 
vessel.     30/1.  SZZI^^I 


ANATOMY   OF  THE   CESTODA 


291 


(fig.  184,  PL) ;  they  lie  internal  to  the  subcuticular  cells  and,  piercing  the 
cuticle  with  their  peripheral  processes,  end  as  projecting  "  receptor  " 
hairs.     Higher  organs  of  sense  are  not  known. 

The  EXCRETORY  APPARATUS  of  the  Cestodes  is  similar  to  that  of 
other  flat  worms.  The  terminal 
(flame)  cells,  which  hardly  differ 
in  appearance  from  those  of 
the  Trematodes,  are  distributed 
throughout  the  parenchyma,  but 
are  more  common  in  the  cor- 
tical than  in  the  medullary  layer 
(fig.  184,  T.c).  Before  opening 
into  a  collecting  tube,  the  capil- 
laries run  straight,  tortuously, 
or  in  convolutions,  anastomos- 
ing frequently  with  one  another 
or  forming  a  retc  niirahile.  The 
collecting  tubes,  which  have 
their  own  epithelial  and  cuti- 
cular  wall,  and  which  also 
appear  to  be  provided  with 
muscular  fibres,  occur  typically 
as  four  canals  passing  through 
the  entire  length  of  the  worm 
(fig.  1 89) ;  they  lie  side  by  side, 
two  (a  wider  thin-walled  ventral, 
and  a  narrower  thick-walled 
dorsal  one)  in  either  lateral  field  ; 
in  the  head  the  two  vessels  on 

each  side  unite  by  means  of  a  loop,  at  the  posterior  extremity  they 
open  into  a  short  pyriform  or  fusiform  terminal  bladder  w^hich  dis- 
charges in  the  middle  of  the  posterior  edge  of  the  original  terminal 
proglottis. 

This  primitive  type  (fig.  189)  of  arrangement  of  collective  tubes  is 
subject  to  variation  in  most  Cestodes,  in  the  scolex  as  well  as  in  the 
segments.  Indeed,  even  the  lumen  of  the  four  longitudinal  tubes 
does  not  remain  equal,  as  the  dorsal  or  external  tubes  are  more  fully 
developed  and  become  thicker,  whereas  the  ventral  or  internal  ones 
remain  thin,  and  in  some  species  quite  disappear  in  the  older  seg- 
ments (figs.  185,  187).  Moreover,  very  frequently  connections  are 
established  between  the  right  and  left  longitudinal  branches,  as  in  the 
head,  where  a  'frontal  anastomosis"  develops,  which  in  the  Tceniidce 
usually  takes  the  form  of  a  ring  encircling  the  rostellum  (fig.  190),  and 
in  the  segments  of  a  transverse  anastomosis  at  each  posterior  border, 


Fkj.  188. — Tcenia  canurus,  head  and  part  of 
neck  showing  nervous  system.  Enlarged.  (After 
Niemiec.) 


292 


THE    ANIMAL   PARASITES   OF   MAN 


especially  between  the  larger  branches,  and  more  rarely  between  the 
smaller  collecting  tubes  also  (fig.  iqO- 

The  so-called  ''  island "  formation  is  another  modification,  i.e., 
at  any  spot  a  vessel  may  divide  and  after  a  longer  or  shorter  course 
the  two  branches  reunite,  and  this  may  appear  in  the  collecting  tubes 
themselves  as  well  as  in  their  anastomoses.  The  above-mentioned  ring 
in  the  frontal  commissure  of  the  Tcemidce  is  such  an  island;  similar 
rings  also  frequently  encircle  the  suckers  (fig.  190).  In  extreme  cases 
[Triceiiophoriis,  Ligiila,  Dihothriocephaliis,  etc.)  this  island  formation 
extends  to  all  the  collecting  tubes  and  their  anastomoses.  Instead  of 
two  or  four  longitudinal  canals  only,  connected  by  transverse  anasto- 
moses at  the  posterior  border  of  the  segments,  there  is  an  irregular 
network   of   vessels,   situated   in    the  cortical    layer,  from  which  the 


Fig.  189. — Young  Acanthobothrium 
coronattivi,  v.  Ben.,  with  the  excretory 
vessels  outlined.  Slightly  enlarged.  (After 
Pintner.) 


Fig.  190. — Scolex  of  a  cysticercoid 
from  Arion  sp.,  with  the  excretory 
vessels  outlined.     (After  Pintner.) 


longitudinal  branches,  having  again  subdivided,  can  only  be  dis- 
tinguished at  intervals,  and  even  then  not  in  their  usual  number. 

The  opening  of  the  longitudinal  branches  at  the  posterior  end 
requires  more  accurate  investigation  ;  it  is  true  that  a  single  terminal 
bladder  is  mentioned  as  being  present  in  many  species,  but  this  is 
also  disputed  ;  when  the  original  end  proglottis  has  been  cast  off,  the 
longitudinal  branches  discharge  separately.  Some  species  possess  the 
so-called  foramina  secundaria,  which  serve  as  outlets  for  the  collecting 
tubes ;  they  are  generally  at  the  neck,  but  may  be  situated  on  the 
segments. 

The  contents  of  the  excretory  vessels  is  a  clear  fluid,  the  regurgi- 
tation of  which  is  prevented  by  the  valves  present  at  the  points  of 
origin  of  the  transverse  anastomoses.  The  fluid  contains  in  solution 
a  substance  similar  to  guanine  and  xanthine. 


ANATOMY  OP  THE  CESTODA 


293 


Genital  Organs. — With  the  exception  of  one  genus  {DioecocestuSy 
Fuhrm.),  in  which  the  species  are  sexually  differentiated,  all  the 
Cestodes  are  hermaphroditic ;  the  genitalia  develop  gradually  in 
the  segments  (never  in  the  scolex),  the  male  organs,  as  is  usual 
in  hermaphroditic  animals,  forming  earlier  than  the  female.  The 
youngest  proglottids  generally  do  not  exhibit  even  traces  of  genitalia  : 
these,  as  a  rule,  develop  first  in  the  older  segments,  and  the  develop- 
ment proceeds  onwards  from  segment  to  segment.  In  a  few  ex- 
ceptional cases  (Lignla)  the  sexual  organs  are  already  developed  in 


Vag-.  Vsc.  Shg. 

Fig.  191. — Proglottis  of  Tania  sagmata,  Goeze,  showing  genitalia.  C,  trans- 
verse excretory  canal ;  N.,  lateral  longitudinal  nerve  ;  IV.,  longitudinal  excretory 
canal ;  7".,  testicles  scattered  throughout  the  proglottis;  Ut.^  opposite  the  central 
uterine  stem  (a  closed  sac)  ;  Ss.^  genital  pore  leading  into  the  genital  sinus  ;  above 
the  cirrus  and  coiled  vas  deferens  {V.d.),  below  the  vagina  [Vag.),  bearing  near 
its  termination  a  dilatation,  the  seminal  receptacle;  Vsc,  the  triangular  vilel- 
larium,  and  above  it  {Shg.)  the  shell  gland  ;  leading  from  this  to  the  uterus  is  seen 
the  short  uterine  canal,  on  either  side  of  this  the  two  lobes  of  the  ovary  {Ov.).  lo/i. 


the  larval  stage,  but  are  only  functional  after  the  entry  of  the  parasite 
into  the  final  host. 

With  the  exception  of  the  end  portions  of  the  vagina,  cirrus  and 
uterus,  all  the  parts  of  the  genital  apparatus  lie  in  the  medullary  layer, 
except  only  the  vitellaria,  which  in  many  species  are  in  the  cortical 
layer.  The  male  apparatus  consists  of  the  testes,  of  which,  as  a  rule, 
there  are  a  large  number,^  and  which  lie  dorsal  to  the  median  plane 
(fig.    185,    T.)  ;    a   vas    efferens    arises    from   each   testis,  unites   with 

'  There  are,  however,  tapeworms  with  only  one,  others  with  only  two  or  three  testes 
in  each  segment. 


294 


THE   ANIMAL   PARASITES   OF   MAN 


contiguous  vasa,  and  finally  discharges  into  the  muscular  vas  deferens 
that  'is  situated  in  about  the  middle  of  the  segment.  According  to 
the  position  of  the  genital  pore,  the  vas  deferens  opens  on  the  lateral 
margin  or  in  the  middle  line  in  the  front  of  the  segment  ;    it  is  much 


Fig.  i^2.—Dibothriocephalus  laius.  Upper  figure  :  female  genitalia,  ventral 
view.  Lower  figure  :  male  genitalia,  dorsal  view.  The  central  portion  only  of 
the  proglottis  is  shown,  a,  cirrus  sac  ;  b,  partly  everted  cirrus  ;  c,  genital  atrium 
and  pore  ;  d,  vaginal  pore  ;  e,  uterus  ;  /,  uterine  pore  ;  g,  vagina  ;  h,  ovary  ; 
i,  shell  gland  ;  j,  vitelline  duct ;  k,  lateral  nerve  ;  /,  vitellarium  ;  n,  vas  deferens 
(muscular  portion)  ;  /,  vas  deferens ;  q,  seminal  vesicle  ;  r  and  x,  vasa  efferentia  ; 
s,  lateral  excretory  canal  ;  /,  testicular  follicles.  (After  Benham  and  Sommer 
and  Landois.) 


ANATOMY   OF   THE   CESTODA 


295 


convoluted  or  twisted,  and  frequently  possesses  a  dilatation  termed 
the  vesicula  seminalis.  It  finally  enters  the  cirrus  pouch,  which  is 
usually  elongated ;  within  the  cirrus  pouch  lies  the  protrusible  cirrus, 
which  is  not  uncommonly  provided  with  hooklets. 

The  male  sexual  orifice  almost  always  opens  with  that  of  the 
vagina  into  a  genital  atrium,  the  raised  border  of  which  rises  above 
the  edge  of  the  segment  and  forms  the  genital  papilla  (fig.  191). 

The  vagina,  like  the  vas  deferens,  usually  runs  inwardly  and 
posteriorly,  where  it  forms  a  spindle-shaped  dilatation  (receptaculum 
seminis) ;     its    continuation,    the    spermatic    duct,    unites    with    the 


Fig.  193. — Diagram  of  genitalia  of  a  Cestode.  g.p.^  genital  pore;  $  ^,  male 
and  female  ducts  opening  into  genital  sinus;  c.s.^  cirrus  sac;  v.d.^  coiled  vas 
deferens  {"  outer  seminal  vesicle") ;  vag.^  vagina  ;  sent,  rec,  seminal  receptacle  ; 
sp.  d.,  spermatic  duct ;  C.c,  fertilization  canal  ;  vit.  d.,  vitelline  duct ;  sk.  g. ,  shell 
gland;  ut.  c,  uterine  canal;  ttt.,  uterus;  Ov.,  ovary;  /,  pumping  organ. 
Cf.  figs.  191  and  233.     (Stephens.) 

oviduct,  the  common  duct  of  the  ovaries  (fig.  191).  The  ovaries, 
usually  two  in  number,  are  compound  tubular  glands  in  the  posterior 
half  of  the  proglottis,  which  extend  into  the  medullary  layer,  but 
ventral  to  the  median  plane. 

At  the  origin  of  the  oviduct  there  is  frequently  a  dilatation  pro- 
vided with  circular  muscles  (suction  apparatus),  which  receives  the 
ovarian  cells  and  propels  them  forward.  After  the  oviduct  has 
received  the  spermatic  duct  the  canal  proceeds  as  the  fertilization 
canal,  and  after  a  very  short  course  receives  the  vitelline  duct  or 
ducts,  and  then  the  numerous  ducts  of  the  shell  glands  (ootype). 
[Although  the  nomenclature  of  these  parts  varies,  we  may  consider 
the  oviduct  as  extending  from  the  ovary  to  the  shell  gland  and 
as  receiving  the  spermatic  duct  and  then  the  vitelline  duct  and  the 
ducts  of  the  shell  gland.  The  short  piece  into  which  the  shell  gland 
ducts  open  corresponds  to  the  ootype  in  the  flukes,  but  in  the  tape- 
worms this  portion  of  the  canal  is  seldom  dilated.     From  this  point 


296 


THE   ANIMAL   PARASITES   OF   MAN 


the  oviduct  is  continued  as  a  shorter  or  longer  tube,  the  uterine  canal 
or  true  oviduct  opening  into  the  uterus  proper. — J.  W.  W.  S.] 
The  vitellariuni  may  be  single,  but  often  exhibits  its  primitive 
duplication  more  or  less  distinctly,  in  which  case  it  is  situated  at  the 
posterior  border  of  the  segments  in  the  medullary  layer  (fig.  191). 
The  original  position  of  the  double  organ  is,  moreover,  the  same 
as  in  the  Trematodes,  i.e.j  at  the  sides  of  the  proglottids,  and  thence 
eventually  extending  more  or  less  on  both  surfaces  (figs.  192  and  194) ; 
the  gland  is  then  distinctly  grape-like  and  the  follicles  lie  mostly  in 
the  cortical  layer. 

The  tgg  cell  that  has  been  fertilized  and  supplied  with  yolk  cells 
receives  the  shell  material  at  the  point  of  entry  of  the  shell  gland  ducts, 
and,  as  a  complete  (^gg,  then  moves  onward  to  the  uterus.  In  those 
cases  in  which  the  uterus  in  its  further  course  presents  a  convoluted 
canal,  and  may  form  a  rosette  (pseudo-phyllidea),  there  is  an  external 


Vvs 


Fig.  194.— Part  of  a  transverse  section  through  a  proglottis  of  Dibothriocephalus  la/us.  Ct.^ 
cuticle;  C,  cirrus;  Vvs.,  vitelline  follicles;  L.M.,  longitudinal  muscles;  T.,  testicles  j 
M.,  medullary  nerve  ;  S.c,  subcuticle  ;    T.m.,  transverse  muscles  ;   UL,  uterus.     20/1. 


opening  which  is  usually  separate  from  the  genital  pore,  and  lies  on 
the  same  or  the  opposite  surface.  In  all  other  cases,  however,  the 
uterus  terminates  blindly  and  is  represented  by  a  longer  or  shorter  sac 
lying  in  the  longitudinal  axis  (fig.  191),  but  in  many  forms  trans- 
versely. With  the  accumulation  of  eggs  it  becomes  modified  in 
various  ways  :  (i)  it  sends  out  lateral  branches  (fig.  241),  or  (2)  forms 
numerous  isolated  sacs  (parenchymal  capsules)  containing  single 
eggs  or  groups  of  eggs  (fig.  217)  ;  further,  (3)  in  some  cases  at  the 
blind   end    one   or   more   special   thick-walled    cavities    are   formed 

(PARUTERINE   ORGANS  Or  UTERINE  CAPSULES),  in  which  all  or  most  of 

the  eggs  are  collected,  the  uterus  then  undergoing  atrophy. 

In  species  in  which  the  uterus  lacks  an  opening,  simultaneously 
with  the  growth  of  this  organ  an  atrophy  of  the  male  apparatus,  at 
least  of  the  testes  and  their  excretory  ducts,  takes  place  ;  this  atrophy 
also  frequently  occurs  in  the  female  glands,  so  that  the  entire  mature 
segments  have  besides  the  uterus  only  traces  of  the  genitalia  left. 


DEVELOPMENT   OF   THE   TAPEWORMS  297 

In  the  Acolei'iicv  the  vagina  is  more  or  less  extensively  atrophied, 
and  in  any  case  has  no  external  opening. 

A  number  of  genera  are  distinguished  by  the  duplication  of 
the  genitalia  in  every  segment ;  the  genital  apparatus  in  its  entirety, 
or  with  the  exception  of  the  uterus,  is  double,  or  the  genital  glands 
and  the  uterus  are  single,  but  the  cirrus,  vas  deferens  and  vagina 
are  double. 

On  comparing  the  genitalia  of  the  Trematodes  and  Cestodes  the 
parts  will  be  found  to  agree,  but  the  vagina  of  the  Cestodes 
corresponds  with  the  uterus  of  the  Trematodes,  and  the  uterus  of  the 
tapeworms  to  Laurer's  canal  of  the  Trematodes,  which  in  most  of  the 
Cestodes  has  lost  its  external  orifice. 

Development  of  the  Tapeworms. 

Copulation. — As  each  proglottis  possesses  its  own  genital  apparatus, 
and  male  as  well  as  female  organs  are  present,  the  following  processes 
may  occur  :  (i)  self-  or  auto-fecundation  (without  immissio  cirri) ; 
(2)  self-  or  auto-copulation  (with  immissio  cirri)  ;  (3)  cross-copula- 
tion between  proglottids  of  the  same  or  different  chains  (of  the  same 
species) ;  and  (4)  cross-copulation  in  the  same  proglottis  in  species 
with  double  genital  pores.  These  various  modes  have  actually  been 
observed. 

In  those  species  w^hich  lack  the  vagina  (Acoletnce)  it  appears  that 
the  cirri,  which  are  always  furnished  with  hooks,  are  driven  into 
the  tissues  and  for  the  most  part  reach  the  receptaculum  seminis. 

The  eggs  of  all  Cestodes  are  provided  with  shells,  but  the  shells, 
like  their  contents,  vary.  In  genera  that  possess  a  uterine  pore  the 
mature  eggs  frequently  do  not  differ  from  those  of  the  Distomata  ; 
they  have  a  brown  or  yellow  shell  of  oval  form  provided  with  an 
operculum,  and  contain  a  number  of  yolk  cells  in  addition  to  the 
fertilized  ovarian  cell  (fig.  128),  but  in  other  genera  (with  a  uterine 
pore)  the  lid  is  absent  and  the  egg-shell  is  very  thin,  the  eggs  of  these 
genera  resembling  those  of  Cestodes  in  which  the  secretion  of  the 
vitellarium  is  a  light  albumin-like  substance  that  contains  only  a 
few  granules,  and  in  which  the  egg-shell  is  very  delicate  and  without 
operculum. 

The  eggs  of  Tmtililce,  for  example,  at  first  consist  of  egg-shell 
(ootype),  ovum  and  yolk  cells.  The  egg-shell  is  as  a  rule  soft,  colour- 
less and  frequently  deciduous,  and  the  yolk  is  scanty  in  amount  and 
contains  few  granules.  The  eggs  are,  moreover,  more  complicated 
than  this.  They  enlarge  and  change  their  shape  and  various  envelopes 
are  developed  around  the  embryo.  The  egg-shell  proper  often  dis- 
appears, and   one    or    more   embryonal    envelopes,    or   protoplasmic 


298 


THE   ANIMAL   PARASITES   OF   MAN 


layers,  arise,  so  that  eventually  it  is  difficult  to  say  whether  the  whole 
egg  is  present,  and,  if  not,  what  the  layers  that  remain  really  are. 

The  embryonal  development  in  most  species  takes  place  during  the 
stay  of  the  eggs  in  the  uterus;  in  other  species  it  takes  place  after  the 
eggs  have  been  deposited  and  are  in  water.  Separate  cells  or  a  layer  of 
cells  always  separate  from  the  segmentation  cells,  as  well  as  from  the 
cells  of  the  developing  embryo,  and  form  one  or  more  envelopes  round 
the  embryo  ;  usually  two  such  envelopes  are  formed,  the  inner  one 
of  which  stands  in  intimate  relationship  with  the  embryo  itself  and 
is  often  erroneously  termed  the  egg-shell,  but  more  correctly  the 
embryonal  shell  or  emhryophore.  In  some  species  it  carries  long  cilia, 
as  in  Dibothriocephalus  latus,  by  aid  of  which  the  young  swim  about 
when  released  from  the  egg-shell  ;  as  a  general  rule,  however,  there 
are  no  cilia  and  this  envelope  is  homogeneous,  or  is  composed  of 
numerous  rods  and  is  calcified,  as  in  Tcenia  spp.  (fig.  197).  The 
second  outer  envelope  ("yolk  envelope  ")  (fig.  207,  3)  lies  close  within 


Fig.  195. — ¥,gg  o{  Diplogono- 
porus  grandis,  showing  the 
morula  surrounded  by  yolk  cells 
and  granules.  440/1.  (After 
Kurimoto.) 


Fig.  196.  —  Uterine  egg  of  Tania  saghiata,  G. 
Uterine  shell  with  filaments  ;  the  oncosphere  with 
embryonal  shell  (embryophore)  in  the  centre. 
500/1.     (After  Leuckart.) 


the  true  (ootype)  egg-shell,  and  remains  within  it  when  the  embryo 
hatches  out,  and  in  many  species,  as  in  Tcenia  spp.,  it  perishes  at  the 
end  of  the  embryonal  development  with  the  delicate  egg-shell  which 
was  formed  in  the  ootype,  so  that  one  observes  not  the  entire  egg 
with  egg-shell  but  only  the  embryo  in  its  embryonal  shell,  viz.,  the 
embryophore  (fig.  197,  a.). 

The  embryo  (the  oncosphere)  enclosed  within  the  embryonal 
shell  (embryophore)  is  of  spheroidal  or  ovoid  form  (fig.  197,  6.),  and  is 
distinguished  by  the  possession  of  three  pairs  of  spines,  a  few  terminal 
(flame)  cells  of  the  excretory  system,  and  muscles  to  move  the  spines. 

No  FURTHER  DEVELOPMENT  of  the  oncosphere  takes  place,  either 
in  the  parent  organism  or  in  the  open  ;  in  fact,  in  all  cases  in  which 
the  oncospheres  are  already  formed  within  the  proglottids  they  do 
not  become  free,  but  remain  in  their  shell  ;  it  is  only  when  the 
oncospheres  are  provided  with  a  ciliated  embryophore  that  they  leave 
the  egg-shell,  and  they  even  cast  this  ciliated  envelope  after  having 


DEVELOPMENT   OF   THE   TAPEWORMS 


299 


swum  about  in  water  by  its  means  for  a  week  or  so.  Sooner  or  later, 
however,  all  the  oncospheres  leave  the  host  that  harbours  the  parental 
tapeworm  and  reach  the  open,  either  still  enclosed  in  the  uterus  of 
the  evacuated  proglottids,  after  the  disintegration  of  which  they 
then  become  free,  or  after  being  deposited  as  eggs  in  the  intestine 
of  the  host;  they  then  leave  it  with  the  faeces.  In  the  former  case 
also,  the  slightest  injury  to  the  mature  proglottids  while  still  in  the 
intestine  suffices  to  allow  a  part  of  the  oncospheres  in  their  embryo- 
phores  to  be  released  and  mingled  with  the  faeces.  Here  they  are  the 
generally,  but  falsely,  so-called  Taeniae  '^  eggs."  For,  as  stated  above, 
the  ''yolk"  envelope  and  the  true  shell  deposited  in  the  ootype  have 
before  this  disintegrated. 

In  other  cases,  e.g.,  Hymenolepis  spp.,    the  uterine  (ootype)  shell 
persists  in  faeces  (fig.  230). 

In  any  case  the  oncospheres 
must  be  transmitted  into  suitable 
animals  to  effect  their  further  de- 
velopment ;  in  only  very  rare  cases 
might  an  active  invasion  be  possible, 
as,  for  instance,  takes  place  with  the 
miracidia  of  many  Trematodes.  The 
entry  into  an  animal  is,  as  a  rule, 
entirely  passive,  that  is  to  say,  the 
oncospheres  are  swallowed  with  the 
food  or  water.  Many  animals  are 
coprophagous  and  ingest  the  onco- 
spheres direct  with  the  faeces  ;  others 
swallow  them  with  water,  mud,  or 
food  contaminated  by  such  faeces, 
artificially  by  feeding  suitable  animals  with  mature  proglottids  of 
certain  Cestodes  or  introducing  the  oncospheres  with  the  food.  As 
the  mature  tapeworm  frequently  finds  the  conditions  suitable  for  its 
development  in  only  one  species  of  host,  or  in  species  nearly  related, 
and  perishes  when  artificially  introduced  into  other  hosts,  experiment 
has  taught  us  that  to  succeed  in  cultivating  the  oncospheres  certain 
species  of  animals  are  necessary.  Thus  we  are  aware  that  the  onco- 
spheres of  Tcvnia  solium,  which  lives  in  the  intestine  of  man,  develop 
only  in  the  pig,  and  only  quite  exceptionally  develop  into  the  stage 
characteristic  of  all  Cestodes — the  cysticercus  in  the  wide  sense  of  the 
word— in  a  few  other  mammals.  The  oncospheres  of  T.  saginaia 
develop  further  only  in  the  ox ;  those  of  T.  marginata  (of  the  dog)  in 
the  pig,  goat,  and  sheep;  those  of  T.  serrata  (of  the  dog)  in  hares  and 
rabbits  ;  those  of  Dipylidiiim  caniniim  (of  the  dog  and  cat)  in  parasitic 
insects  of  the  dog  and  cat,  etc.     It  is  not  unusual  that  young  animals 


Fig.  197. — a.,  oncosphere,  in  its 
radially  striated  embryophore  (errone- 
ously termed  egg-shell)  of  Tania 
africana.  Greatly  magnified.  (After 
von  Linstow.)  b.,  freed  oncosphere  of 
Dipylidium  caninum.  (Alter  Grassi 
and  Rovelli.)  Both  oncospheres  show 
six  spines. 


Infection    is    easily    produced 


300  THE   ANIMAL   PARASITES   OF   MAN 

only  appear  to  be  capable  of  infection,  while  older  animals  of  the 
same  species  are  not  so. 

Once  introduced  into  a  suitable  animal,  which  is  only  excep- 
tionally the  same  individual  or  belongs  to  the  same  species  as  the 
one  which  harbours  the  adult  tapeworm,  the  oncosphere  passes 
into  the  larval  stage  common  to  all  Cestodes,  but  varying  in  structure 
according  to  the  species.  In  the  simplest  case — as,  e.g.,  in  Dibothrio- 
cephahis — such  a  larva  resembles  the  scolex  of  the  corresponding 
tapeworm,  only  that  the  head,  provided  with  suckers,  is  retracted 
within  the  fore-part  of  the  neck.  Such  a  larval  form  is  known  as  a 
plerocercoid  {ifKrjpT]^,  full;  KepKo^,  tail).  They  differ  from  the  cysticer- 
coids  in  being  solid  larval  forms,  elongated,  tape-like  or  oval,  with  the 
head  invaginated.  The  conditions  appear  to  be  similar  in  Ligiila, 
Schistocephalns,  Tricenopliorns,  but  here  the  larvae  are  very  large,  indeed 
as  large  iii  the  first-mentioned  genera  as  the  tapeworms  originating 
from  them,  and  the  sexual  organs  are  already  outlined ;  doubtless, 
however,  this  stage  is  preceded  by  one  that  corresponds  to  the  scolex 
of  the  genus  in  question,  and  which  represents  the  actual  larval  stage. 
In  such  cases  the  development  of  the  body  of  the  tapeworm  from  the 
scolex  has  already  begun  within  the  first  or  intermediate  host ;  in 
other  cases,  except  in  the  single-jointed  (monozootic)  Cestodes,  this 
only  takes  place  in  the  definitive  host.  The  direct  metamorphosis 
of  the  oncosphere  into  the  larval  forms  termed  PLEROCERCOID  has 
hitherto  not  been  investigated,  although  Ligiila,  SchlstocepJialus  and 
Bothriocepliahis  are  very  common  parasites,  but  many  circumstances 
point  to  the  conclusions  arrived  at  by  us  and  by  other  observers.  In 
the  larval  stages  of  other  tapeworms  we  can  always  distinguish  the 
scolex  and  a  caudal- like  appendage,  vesicular  in  the  cysticerci  (fig.  200), 
compact  in  the  cysticercoids  (fig.  231).  The  scolex  alone  forms  the 
future  tapeworm,  the  variously  formed  appendage  perishing. 

It  has  now  been  proved  that  the  appendage,  the  caudal  vesicle, 
originates  direct  from  the  body  of  the  oncosphere,  and  therefore  is 
primary,  and  that  the  scolex  only  subsequently  forms  through  pro- 
liferation on  the  surface  of  this  appendage.  On  account  of  this 
origin  the  scolex  is  generally  regarded  as  the  daughter,  and  the  part 
usually  designated  as  the  appendage  as  the  mother,  originating  from 
the  oncosphere. 

Accordingly,  two  modes  of  development  of  the  larval  stage  may 
be  distinguished  ;  in  the  one  case,  plerocerci  and  plerocercoids,  the 
oncosphere  changes  directly  into  the  scolex,  thus  forming  the  body 
of  the  tapeworm  within  the  primary  host ;  in  the  other  case,  cysticerci 
and  cysticercoids,  the  scolex  only  forms  secondarily  in  the  trans- 
formed body  of  the  oncosphere,  which  later  on  perishes,  the  scolex 
alone  remaining  as  the  originator  of  the  tapeworm  colony. 


DEVELOPMENT   OF   THE   TAPEWORMS 


301 


We  may  summarize  briefly  what  has  been  said  regarding  these 
larval  forms.  We  have,  firstly,  sohd  larval  forms  without  any  bladder. 
These  arise  directly  from  the  oncosphere  and  are  of  two  kinds, 
plerocercus  and  plerocercoid.  Plerocercus  is  a  solid  globular  larva  with 
the  head  invaginated  into  the  posterior  portion.  Plerocercoid  (fig.  208) 
is  a  solid  elongated  larva  also  with  the  head  invaginated  mto  the 
posterior  portion,  which  is  sometimes  very  long.  Secondly,  we 
have  larval  forms  with  bladders  from  which  the  scolices  arise  thus 
indirectly  from  tlie  oncosphere.  They  are  of  two  kinds,  cysticercoid 
and  cysticercus. 

Cysticercoid. — The  bladder  is  but  slighUy  developed  and  is  usually 
absorbed  again.  The  anterior  portion  is,  moreover,  retracted  into 
the  posterior,  and  in  some  cases  there  is  a  long  or  a  stumpy  tail 
(figs.  220,  231). 

Cysticercns,  or  true  bladder  worms.  (These  may  be  divided  into 
[i]    cysticercus    proper,  consisting   of    a    bladder    and    one   scolex ; 


sec.c 


c.v. 


Fig.  198. — Diagram  of  a  cysticercoid.  Cf. 
figs.  220,  227.  c.v.y  caudal  vesicle  or  bladder 
(small)  ;  sec.  c,  secondary  cavity  caused  by  the 
growth  forward  of  the  hind-body;  /.,tail  bearing 
six  spines.     (Stephens.) 


C.V. 


Fig.  199. — Diagram  of  a  cysticercus. 
C.V.,  caudal  vesicle  or  bladder;  i.,  in- 
vagination of  wall  of  bladder.    (Stephens.) 


[2]  coenurus,  a  bladder  and  many  scolices  ;  [3]  echinococcus,  a 
bladder  in  which  daughter  bladders  or  cysts  are  developed,  and  then 
in  these  multiple  scolices.) 

In  the  case  of  cysticerci  a  papilliform  invagination  forms,  pro- 
jecting into  the  interior  of  the  bladder  (fig.  201).  The  layer  of  cells 
forming  the  papilla  becomes  divided  into  two  laminae,  the  outer^  of 
which  forms  a  kind  of  investing  membrane  (receptaciilum  capitis)  for 
the  papilla.  The  head  and  suckers  are  now  developed  on  the  walls 
bounding  the  axial    lumen    of   the    papilla.      The  papilla  eventually 


I.e.,  regarded  from  the  interior  or  centre  of  the  invagination. 


302  THE   ANIMAL   PARASITES   OP^   MAN 

evaginates,  so  that  the  receptaculum  capitis  now  forms  the  inner 
surface  of  the  hollow  head,  which  eventually  becomes  solid. 

Our  knowledge  of  the  development  of  cysticerci  in  the  wide 
sense  of  the  word  is  limited  almost  exclusively  to  that  of  a  few  true 
'^bladder  worms"  (cysticerci);  in  other  cases  we  know  either  only 
the  terminal  stage,  i.e.j  the  complete  larva,  or,  exceptionally,  one  of 
the  intermediate  stages,  but  we  are  not  acquainted  with  a  complete 
series ;  the  description  must  therefore  be  incomplete. 

We  know  from  feeding  experiments  that,  after  the  introduction 
of  mature  proglottids  or  of  the  fully  developed  ova  of  Tcenia 
crassicollis  (of  the  cat)  into  the  stomach  of  mice,  the  oncospheres 
escape  from  the  shell  in  the  middle  portion  of  the  small  intestine, 
and  a  few  hours  later  penetrate  into  the  intestinal  wall  by  means 
of  a  boring  movement ;  they  have  been  found  in  this  position  twenty- 
seven  to  thirty  hours  after  the  infection.  By  means  of  this  migration, 
for  which  purpose  they  employ  their  spines,  they  attain  tke  blood- 
vessels of  the  intestine;  indeed,  already  nine  hours  after  the  infection 
and  later  they  are  found  in  the  blood  of  the  portal  vein,  and  in  the 
course  of  the  second  day  after  infection  they  are  found  in  the 
capillaries  of  the  liver,  which  these  larvae  do  not  leave. 

Leuckart,  in  experimental  feeding  of  rabbits  with  oncospheres 
of  Tcenia  serrata  (of  the  dog),  found  free  oncospheres  in  the  stomach 
of  the  experimental  animal,  but  not  in  the  intestine  :  however, 
he  came  across  them  again  in  the  blood  of  the  portal  vein.  The 
passage  through  the  blood-vessels  to  the  liver  is  the  normal  one 
for  those  species  of  Tcvnia  the  eggs  of  which  become  larvae  in 
mammals ;  even  in  those  cases  in  which  the  oncospheres  develop 
further  in  the  omentum  or  in  the  abdominal  cavity  {Cysticercits 
temiicollis,  C.  pisifonnis),  there  are  distinct  changes  observable  in 
the  liver  that  lead  one  to  the  conclusion  that  there  has  been  a 
secondary  migration  out  of  the  liver  into  the  abdominal  cavity. 
Indeed,  one  must  not  imagine  that  the  young  stages  of  the  Cestodes 
are  absolutely  passive  ;  once  they  have  invaded  an  organ  they  travel 
actively,  and  leave  distinct  traces  of  their  passage. 

In  other  cases  the  oncospheres  leave  the  liver  with  the  circulation, 
and  are  thus  distributed  further  in  the  body;  they  may  settle  and 
develop  in  one  or  more  organs  or  tissues.  Many  oncospheres  may, 
by  travelling  through  the  intestinal  wall,  penetrate  through  it  and 
attain  the  abdominal  cavity  direct  ;  some,  perhaps,  pass  also  into  the 
lymph  stream.  Where  there  are  no  blood  and  lymphatic  vessels  in 
the  intestinal  wall,  as  in  insects,  the  oncospheres  attain  the  body 
cavity  or  its  organs  direct  ;  in  short,  they  never  remain  in  the 
intestinal  lumen  itself,  and  only  rarely — as  in  Hyrnenolepis  murina 
of  the  rat — do  they  remain  in  the  intestinal  wall. 


DEVELOPMENT   OF   THE   TAPEWORMS 


303 


When  the  infection  has  been  intense,  and  the  body  is  crowded  with  numerous 
oncospheres,  acute  feverish  symptoms^  are  induced,  to  which  the  infected  animals 
usually  succumb  ("  acute  cestode  tuberculosis  ")  ;  while  in  other  cases  the  alterations 
in  the  organs  attacked— as  the  liver  in  mice  and  the  brain  in  sheep — may  cause  death. 

Sooner  or  later  the  oncospheres  of  tapeworms  come  to  rest, 
and  are  first  transformed  into  a  bladder,  which  may  be  round  or 
oval  according  to  the  species.  The  embryonal  spines  disappear  sooner 
or  later,  or  remain  close  together  or  spread  over  some  part  of  the 
bladder  wall  (fig.  200).  Their  discovery  by  v.  Stein  in  the  bladder 
worm  of  the  ^'  meal  \vorm  "  (the  larva  of  a  beetle,  Tenebrio  molitor) 
first  led  to  the  conclusion  that  bladder  worms  (cysticerci)  actually 
originate  from  the  oncospheres  of  Tceniidcv. 


Fig.  200. — Diagram  of  dtvelopment  of  a  cysticercus.  i,  solid  oncosphere  with  six  spines  ; 
2,  bladder  formed  by  liquefaction  of  contents  ;  3,  invagination  of  bladder  wall ;  4,  formation 
of  rostellum  (with  booklets)  and  suckers -at  the  bottom  of  the  invagination  ;  5,  evagination  of 
head;  6,  complete  evagination  effected  by  pressure.     (Stephens.) 

The  bladder  may  remain  as  a  bladder,  and  then  by  proliferation 
the  .scolex  forms  on  its  wall  (fig.  202),  or  it  may  divide  into  an 
anterior  so-called  "cystic"  portion  and  a  solid  tail-like  appendage  of 
various  lengths,  on  which  the  embryonal  hooks  are  to  be  found, 
and  this  is  particularly  the  case  in  those  larval  forms  (cysticercoids), 
e.^.,  those  of  Dipylidhun  canimim,  that  develop  in  invertebrate  animals, 
such  as  Arthropoda. 

As  mentioned  above  one  may  regard  the  scolex  as  an  individual 
that  originates  through  proliferation  of  the  wall  of  the  parent  cyst, 
mostly  singly,  but  in  those  cysticerci  that  are  termed  coenurus 
(fig.  201)  many  scolices  occur,  whereas  in  those  called  echinococcus 
the  parent  cyst  originating  from  the  oncosphere  of  Tceiiia  echinococcus 
(of  the  dog)  first  produces  a  number  of  daughter  cysts,  which  in  their 


3G4 


THE    ANIMAL   PARASITES   OF   MAN 


turn  form  numerous  scolices.  Echinococcus-like  conditions  also 
occur  in  cysticercoids,  as,  for  instance,  in  those  peculiar  to  earth- 
worms ;  and  similar  conditions  prevail  in  a  larval  form  known  as 
Staphylocystis,  found  in  the  wood-louse  (Gloineris).  Thus  it  happens  in 
these    cases    that    finally  one   tapeworm    egg  produces  not  one,  but 

numerous  tapeworms,  for,  under 
favourable  conditions,  each  scolex 
can  form  a  tapeworm. 


Fig.  201.  — Section  through  a 
piece  of  a  Ccemiriis  cei'ebralis, 
with  four  cephalic  invaginations 
in  different  stages  of  develop- 
ment. At  the  bottom  of  the 
invaginations  the  rostellum,  hooks 
and  suckers  develop.  (From  a 
wax  model.) 


Fig.  202. — Median  section  through 
a  cysticercus,  with  developed  scolex 
at  the  bottom  of  the  invagination. 
(After  Leuckait.) 


The  rudiment  of  the  scolex  appears  as  a  hollow  bud,  the  cephalic  invagination 
usually  directed  towards  the  interior  of  the  bladder  cavity;  on  its  invaginated 
surface  arise  the  four  suckers,  and  the  rostellum  with  the 
hook  apparatus  is  formed  in  its  blind  end  ;  we  thus  get  a 
T.ienia  head,  but  with  the  position  of  the  parts  reversed 
(fig.  201).  In  many  cysticerci  the  head  rises  up  from  the 
base  of  the  cephalic  invagination  and  is  then  surrounded  by 
the  latter.  A  more  or  less  elongated  piece  of  neck  also 
develops,  and  even  proglottids  may  appear,  as  in  Cysticercus 
\  fasciolaris  (the  larva  of  Tcpnia  crassicolUs  of  the  cat)  of  the 
Muridas,  a  process  somewhat  analogous  to  that  of  Ligula,  etc. 

The  period  that  elapses  from  the  time  of 
.^  infection  till  the  cysticercus  is  fully  developed 
varies  according  to  the  species  ;  the  cysticercus 
of  Tcvnia  saginata  requires  twenty-eight  weeks, 
that  of  T.  marginaia  seven  to  eight  weeks,  that 
of  T.  solium  three  to  four  months,  and  that  of 
T.  echinococcns  longer  still. 

With  one  single  exception  {Archigetes)  the 
larvae  do  not  become  sexually  mature  in  the  organ 
where  they  have  developed  ;  they  must  enter  the 
terminal  host,  a  matter  that  is  usually  purely 
passive,  the  carriers  of  the  larvae  or  infected  parts 
of  them  being  usually  devoured  by  other  animals.  In  this  manner,  for 
instance,  the  larvae  (Cysticercns  fasciolaris)  found  in  mice  and  rats  reach 


V 


Fig.  ^wj — Cysticercus 
pisifor?nis  in  an  evagi- 
nated  condition,  with 
neck,  fore-body  and 
bladder,  with  excretory 
network  in  its  wall.  18/1. 


DEVELOPMENT   OF  THE   TAPEWORMS  305 

the  intestine  of  cats  ;    those  of  the  hare  and  rabbit  (C.  pisiformis)  reach 
the  intestine  of  dogs;  those  of  the  pig  (C.  celluloses)  are  introduced  into 
man  ;  those  of  insects  are  swallowed  by  insectivorous  birds ;  those  of 
crustaceans  are  ingested  by  ducks  and  other  water  fowl  ;    perhaps, 
also,   the    infection    of    herbivorous    mammals    is    caused    by   their 
accidentally  swallowing  smaller  creatures  infected  by  larvae.     Indeed, 
the   researches  of   Grassi  and    Rovelli    have   taught  us  that  such  an 
intermediate  host  is   not   always  necessary ;    Hymenolepis  nmrina  of 
rats  and  mice  in  its  larval  stage  lives  in  the  intestinal  wall  of  these 
rodents,  and  as  a  larva  it  passes  into  the  intestinal  lumen  and  develops 
into  a  tapeworm  in  exactly  the  same  way  as  the  larvae  of  other  species 
that  reach  the  intestine  of  the  terminal  host  by  means  of  an  inter- 
mediate carrier.     Probably  this  curtailed  manner  of  transmission  also 
occurs  in  many  other  species.     In  some  cases  the  larvae  actively  quit 
the  body  of  the  intermediate  host,  as  in  the  case  of  Ligula  and  Schisto- 
cepJiahis,  which  travel    out   of   the  body  cavity  of  infected  fish  and 
reach  the  water,  where  they  may  be  observed  in  hundreds  in  summer, 
at  all  events  in  some  localities.     The  larval  stage  of  Calliobothriuin — 
wrongly  termed  Scolex — has  been  observed  swimming  free  in  the  sea, 
and  the  scolices  of  Rhynchobolhriiun,  without  their  mother  cysts,  have 
been  observed  free  within  the  tissues  of  several  marine  animals.     In 
any  case  there  is  almost  always  a  change  of  hosts,  even  in  the  single- 
jointed  Cestodes,  for  the  larva  of  Caryophyllceus,  which  lives  in  fishes 
of  the  carp  family,  is  found  in  limicoline  Oligochaetes,  that  of  Gyro- 
cotyle  (Chimaera)  in  shell-fish  (Mactra),  and  different  conditions  can 
hardly  be  possible  for  Amphilina.     Archigetes  alone  becomes  sexually 
mature  in  the  larval  stage,  but  the  life-history  of  this  creature  is  not 
well  known,  so  that  it  is  not  impossible  that  the  attainment  of  sexual 
maturity  as  a  larva  in  invertebrates   (Oligochaetes)  is  perhaps  abnor- 
mal, and   somewhat   analogous   to   the   maturity   of   some  encysted 
Trematodes. 

The  METAMORPHOSIS  OF  THE  LARVA  into  the  tapeworm  is  rarely 
accomplished  in  a  simple  manner  ;  the  transformation,  however,  is 
not  complex  in  the  single-jointed  Cestodes,  nor  in  Ligula  and  Schisto- 
cephalus ;  the  latter  is  swallowed  by  birds  (Mergus,  Anas,  etc.),  produces 
eggs  after  only  a  few  days,  and  very  soon  quits  the  intestine  of  its 
terminal  host.  In  all  other  cases  it  is  the  scolex  only  which,  by 
proliferating  at  its  posterior  extremity,  forms  the  proglottids,  after 
having  invaded  as  a  larva  the  intestine  of  a  suitable  host.  The  mother 
cysts,  or  what  corresponds  to  them,  die,  are  digested,  absorbed,  or 
perhaps  even  eliminated  ;  on  the  contrary,  segments  found  on  the 
scolex  during  the  larval  stage,  also  in  the  case  of  Cysticercus  fasciolaris, 
are  retained.  It  is  not  certain  whether  the  larvae  of  Dibothriocephalus 
lose  any  part. 
20 


306  THE   ANIMAL   PARASITES   OF   MAN 

The  time  required  by  the  scolex  to  complete  the  entire  chain 
of  proglottids  does  not  depend  only  on  the  number  it  has  to 
produce,  for  Taenia  echinococaiSy  which,  as  a  rule,  only  possesses 
three  or  four  segments,  takes  quite  as  long  a  time  for  their  growth 
(eleven  to  twelve  weeks)  as  T.  solium  with  its  numerous  seg- 
ments ;  T.  ccenitrus  is  fully  developed  in  three  to  four  weeks,  and 
the  same  holds  good  for  DihotJirlocephalus  latus,  which  possesses 
many  more  segments  than  the  above-mentioned  Taenia  of  th^  -dog. 
In  a  number  of  species  it  has  been  possible  to  determine  fairly 
accurately  the  average  daily  growth  ;  for  instance,  in  Dibothriocephalus 
lattis  the  daily  growth  is  8  cm.,  in  Tcvnia  saginaia  7  cm.,  etc. 

The  history  of  the  development  of  the  Cestodes  demonstrates  that 
persons  and  beasts  harbouring  larval  tapeworms  have  become 
infected  by  having  swallowed  the  oncospheres  of  the  species  of 
tapeworm  to  which  they  belong.  In  regard  to  Hymenolepis  mnrina 
alone,  it  is  known  that  the  introduction  of  the  oncospheres  into  those 
species  of  animals  which  harbour  the  adult  tapeworm  leads  to  the 
formation  of  the  latter  after  the  development  of  a  larval  stage  in  the 
intestinal  w^all ;  nevertheless,  only  young  animals  (rats)  are  capable  of 
infection,  for  a  previous  infection,  or  the  presence  of  mature  tape- 
worms in  the  intestine,  appears  to  produce  a  kind  of  immunity. 

Biology. 

In  their  adult  stage,  the  tapeworms  inhabit  almost  exclusively 
the  alimentary  canal  of  vertebrate  animals,  with  but  few  exceptions 
the  small  intestine,  and  a  few  species  select  definite  parts  of  it.  A 
small  number  of  Rhynchohothriidce  of  marine  fishes  live  apparently 
always  in  the  stomach,  while  in  rays  and  sharks  the  spiral  intestine  is 
their  exclusive  site.  Bothriocephali  generally  attach  themselves  with 
their  head  on  to  the  appendices  of  the  pylorus  of  fishes  ;  other 
species  (Hymenolepis  diminuta)  occasionally  fix  their  head  in  the 
ductus  choledochus,  and  this  is  more  frequent  still  in  the  tapeworms 
of  the  rock  badger  (Hyrax),  which  occasionally  penetrate  entirely 
into  the  biliary  ducts.  Stilesia  hepatica,  Wolffh.,  has  so  far  only  been 
found  in  the  bile-ducts  of  its  host  (sheep  and  goat,  East  Africa). 

In  the  disease  of  sheep  induced  by  Cestodes,  the  w^orms  have 
been  observed  also  in  the  pancreas.  Specimens  found  in  the  large 
intestines  were  probably  being  evacuated. 

The  Cestodes  are  looked  upon  as  fairly  inert  creatures,  this 
opinion  having  been  formed  by  observing  their  condition  in  the  cold 
cadavers  of  warm-blooded  animals.  Actually,  however,  they  are 
exceedingly  active,  and  accomplish  local  movements  within  the 
intestine,  for  they  have  been  found  in  the  ducts  communicating  with 


BIOLOGY   OF  THE   CESTODES  307 

the  bowel,  or  in  the  stomach,  and  may  even  make  their  way  forward 
into  the  oesophagus. 

They  also  invade  other  abdominal  organs  through  abnormal  com- 
munications, or  through  any  that  may  be  temporarily  open  between 
the  intestine  and  such  organs  ;  they  thus  reach  the  abdominal  cavity  or 
the  urinary  bladder,  or  they  work  their  way  through  the  peritoneum. 

They  produce  changes  in  the  intestinal  mucous  membrane  at  the 
place  of  their  attachment,  the  alterations  varying  in  intensity  accord- 
ing to  the  structure  of  the  fixation  organs.  The  mucous  membrane 
is  elevated  in  knob-like  areas  by  the  suckers  ;  the  epithelial  cells 
become  atrophied  or  may  be  entirely  obliterated.  Dipylidium 
caninnni  bores  into  the  openings  of  Lieberkiihn's  glands  with  its 
rostellum,  dilating  the  lumen  to  two  or  three  times  its  normal  size, 
while  the  suckers  remain  fixed  between  the  basal  parts  of  the  cells. 
Species  with  powerful  armatures  penetrate  deeper  into  the  submucosa, 
and  some  that  are  not  provided  with  exceptionally  strong  armatures, 
or  are  even  unarmed,  may  be  actually  found  with  the  scolex  embedded 
in  the  muscles  of  the  intestinal  walls  or  even  protruding  beyond 
{Tcenia  tetragonal  MoL,  in  fowls,  etc.).  Other  species,  again,  even 
cause  perforation  of  the  walls  of  the  intestine  of  their  hosts. 

It  is  generally  assumed  that  tapeworms,  which  almost  without 
exception  live  in  the  gut  of  vertebrates,  get  their  nutriment  from  the 
gut  contents,  which  apparently  they  absorb  through  the  whole  body 
surface  (cuticular  trophopores).  In  favour  of  this  view  is  the 
existence  of  fat  drops  in  the  proglottids,  the  identity  in  colour  in 
certain  forms  between  that  of  the  fresh  worm  and  the  gut  contents 
and  the  passage  of  certain  substances  derived  from  medicines  (iron 
and  mercury  preparation)  into  the  worms  in  the  gut,  etc.  Whether 
the  suckers  are  concerned  in  the  absorption  of  nutriment  and  to 
what  extent  is  still  questionable. 

The  length  of  life  of  the  adult  tapeworm  certainly  varies  ; 
as  a  rule  it  appears  to  last  only  about  a  year ;  in  other  cases  (Ligula) 
it  averages  only  a  few  days,  but  we  are  likewise  aware  that  certain 
species  of  Cestodes  of  man  attain  an  age  of  several  or  many  years 
(thirty-five).  The  natural  death  of  Cestodes  often  appears  to  be 
brought  about  by  alterations  in  the  scolex,  such  as  loss  of  the  hooks, 
atrophy  of  the  suckers  and  rostellum,  finally  the  dropping  off  of  the 
scolex;  it  is  unknown  whether  a  chain  of  segments  deprived  of  its 
scolex  then  perishes  or  whether  it  first  attains  maturity.  It  has  already 
been  mentioned  that  in  a  few  species  the  foremost  proglottids  are 
transformed  into  organs  of  fixation  on  the  normal  loss  of  the  scolex. 

Abnormalities  and  malformations  are  encountered  relatively  frequently  in  the 
Cestodes — such  as  abnormally  short  or  long  segments  ;  the  so-called  triangular 
tapeworms,  which — if  belonging  to  the  Tceniidce — always  possess  six  suckers  ;  often 


308  THE   ANIMAL   PARASITES   OF   MAN 

also  club-shaped  segments  occur  between  normal  ones,  or  there  may  be  a  defect 
in  one  segment  or  in  the  centre  of  a  number  following  one  another  (fenestrated 
segments)  ;  bifurcated  chains  of  segments  have  likewise  been  observed,  as  well  as 
incomplete  or  complete  union  of  the  proglottids,  abnormal  increase  of  the  genital 
pores,  reversion  of  the  genitalia.  Besides  the  above-mentioned  increase  of  the 
number  of  suckers  on  the  scolex  (in  Taeniae),  there  may  be  a  decrease  in  the 
number ;  in  other  cases  the  crown  of  hooks  may  be  absent,  or  abnormally  shaped 
hooks  may  be  formed. 


Classification  of  the  Cestoda  of  Man. 
Order.     Pseudophyllidea,  Cams,  1863. 

Scolex  without  proboscis  or  rostellum.     Head  "  stalk"  absent. 

Scolex  never  with  four,  generally  with  two  (or  one  terminal)  bothria.^  Vitellaria 
numerous.  Uterine  opening  present.  Genitalia  do  not  atrophy  when  uterus  is 
developed.  In  large  majority  of  proglottids  eggs  (or,  if  formed,  their  contents)  are 
at  the  same  stage  of  development. 

Family.     Dibothriocephalidae,  Liihe,  1Q02. 

Syn. :  Diphyllobothritdce,  Liihe,  1910. 

Genitalia  repeated  in  each  proglottid  (polyzootic  Cestodes).  Ventral  and  dorsal 
surfaces  flat.  Cirrus  unarmed.  Cirrus  and  vagina  if  non-marginal  open  on  the 
same  surface  as  the  uterus.  Uterus  long,  convoluted,  often  forming  a  "  rosette,"" 
never  dilates  into  a  uterine  cavity.  Eggs  thick  shelled,  operculated,  constantly 
being  formed  in  mature  proglottids. 

Sub-family.     Dibothriocephalinae,  Liihe,  1899. 

Syn.  :    Diphyllobothriina^  Liihe,    19 10. 

Segmentation  distinct.  Scolex  unarmed,  elongated,  sharply  separated  (generally 
by  a  neck)  from  the  first  proglottis.  Cirrus  and  vagina  open  ventrally.  Genital 
pores  non-alternating.  Vas  deferens  surrounded  by  a  muscular  bulb.  Receptacu- 
lum  seminis  large,  sharply  separated  from  the  spermatic  duct. 

Order.     Cyclophyllidea,  v.  Beneden. 

Four  suckers  always  present.  Uterine  opening  absent.  Vitellarium  single. 
Genitalia  atrophy  when  uterus  is  fully  developed. 


'  Bothridia  or  '' phyllidia  "  are  outgrowths  from  the  scolex.  They  are  concave  and 
extremely  mobile.  By  some  authors  the  term  '' phylliditim  "  is  used  for  the  outgrowth,  and 
the  term  '' bothridmm"  is  restricted  to  the  muscular  cup.  Bothria,  on  the  other  hand,  are 
grooves  more  or  less  wide,  the  musculature  of  which  is  only  slightly  developed  and  is  not 
separated  off  internally  from  the  parenchyma.  Acetabula,  or  suckers  in  the  usual  sense,  are 
hemispherical  cups,  without  lips  and  with  musculature  separated  internally  from  the 
parenchyma. 


CLASSIFICATION   OF  THE   CESTODA  309 

Family.     Dipylidiidae,  Liihe,  1910. 

Rostellum  if  present  armed.  Suckers  unarmed.  Uterus  breaks  up  into  egg 
capsules.     Paruterine  organs  absent. 

Family.     Hymenolepididae,  Railliet  and  Henry,  1909. 

Segment  always  broader  than  long.  Genitalia  single.  Longitudinal  muscles  in 
two  layers.  Genital  pores  unilateral.  Testes  one  to  four.  Uterus  persistent,  sac- 
like.    Eggs  with  three  shells. 

Family.     Davaineidae,  Fuhrmann,  1907. 

Rostellum  cushion-shaped.  Armed  with  numerous  (sixty  to  several  thousand) 
hammer-shaped  hooks  in  two  (rarely  one)  rows. 

Sub-family.     Davainelnae,  Braun,  1900. 
Suckers  armed.     Uterus  breaks  up  into  egg  capsules.     Paruterine  organs  absent. 


Family.     Taeniidae,  Ludwig,  1886. 

Suckers  unarmed.  Uterus  with  median  longitudinal  stem  and  lateral  branches. 
Female  genitalia  at  the  hind  end  of  the  proglottis.  Genital  pore  irregularly  alter- 
nating. Testes  numerous  in  front  of  female  genitalia.  Ovary  with  two  lobes  (wings). 
Vitellarium  behind  the  ovary.     Embryophore  radially  striated. 


The  Cestodes  of  Man. 

Most  of  the  species  to  be  mentioned  live  in  man  in  their  adult 
stage  and  occupy  the  small  intestine ;  man  is  the  definite  host  of  these 
parasites,  but  is  not  the  specific  host  for  all  the  species  ;  some  of  these 
-species,  as  well  as  others  (of  mammals),  may  occur  in  man  also  in  the 
larval  stage. 

Family.     Dibothrlocephalldae. 

Sub-family.     Dibothriocephallnae. 

Genus.      Dibothriocephalus,  Liihe,  1899. 

Syn. :   Diphyllobothriiun^  Cobbold,  1858;   Bothriocephalus,  ^.  ^.  Rvid.,  18 19; 
Dibothrius^  p.  p.  Rud.,  1819  ;  Dibothriimi,  p.  p.  Dies.,  1850. 

Scolex  egg-shaped  ;  dorsal  and  ventral  bothria  elongated,  moderately  strong, 
-cutting  rather  deeply  into  the  head  ;  genitalia  single  in  each  proglottis  ;  papillae  in 
the  vicinity  of  the  genital  atrium  ;  the  testes  and  vitellaria  are  in  the  lateral  fields,  the 


310  THE   ANIMAL   PARASITES   OF    MAN 

former  in  the  medullary  layer,  the  latter  In  the  cortical  layer  on  both  surfaces,  and 
occasionally  extending  to  the  median  line  ;  the  ovary  ventral,  the  shell  gland  dorsaL 
The  uterus  is  in  the  central  field,  taking  a  zigzag  course,  and  frequently  forms  a 
rosette. 


Dlbothriocephalus  latus,  L.,  1748. 

Syn. :  TcBtiia  lata,  L.,  1748;  Tcenia  vulgaris,  L.,  1748;  Tcenia  grisea,  Pallas, 
1796;  Tcenia  ineinbrajiacea.  Pall.,  1781  ;  Tania  tenella,  Pall.,  1781  ;  Tcenia  dentatay 
Batsch,  1786;  Boihriocephalus  latus,  Bremser,  1819;  Dibothrium  latinn.  Dies., 
1850;  Boihriocephalus  cristatus,  Davaine,  1874'  ;  Boihriocephalus  balticusy 
Kchnmstr.,  1855;  Boihriocephalus  latissimus,  Bugn.,  1886. 

Length  2  to  9  m.  or  more ;  colour  yellowish-grey ;  after  lying  in 
water  the  lateral  areas  become  brownish  and  the  uterine  rosette 
brown.  The  head  is  almond-shaped,  2  to  3  mm.  in  length,  the  dorso- 
ventral  axis  is  longer  than  the  transverse  diameter ;  the  head,  therefore^ 
generally  lying  flat,  conceals  the  suctorial  grooves  at  the  borders ; 
these  suckers  are  deep  and  have  sharp  edges  (fig.  205).  The  neck 
varies  in  length  according  to  the  degree  of  contraction  and  is  very 
thin  ;  there  are  3,000  to  4,200  proglottids  and  there  may  be  more  ; 
their  breadth  is  usually  greater  than  their  length,  but  in  the  posterior 
third  of  the  body  they  are  almost  square,  and  the  very  oldest  are  not 
uncommonly  longer  than  they  are  broad.  There  are  numerous  testes 
situated  dorsally  in  the  medullary  layer  of  the  lateral  fields ;  the  vas 
deferens  (fig.  192)  passes  dorsally  in  transverse  loops  in  the  central 
field  anteriorly  and  forms  a  seminal  vesicle  before  its  entry  into  the 
large  cirrus  pouch. 

The  orifice  of  the  vagina  is  close  behind  the  orifice  of  the  cirrus  ; 
the  former  passes  almost  straight  along  the  median  line  posteriorly,. 
and  widens  into  a  receptaculum  seminis  shortly  before  its  junction 
with  the  oviduct ;  the  ovary  is  bilobed,  in  shape  like  the  wings 
of  a  butterfly,  ventrally  in  the  medullary  layer ;  the  shell  glands  lie 
in  the  posterior  recess  of  the  ovary ;  the  uterus,  forming  numerous 
transverse  convolutions,  passes  ventral  to  the  vas  deferens  forwards. 
Eggs  (fig.  207)  large,  with  brownish  shells  and  small  lids,  68  /a  to  71/1 
by  45  A*-;  the  ovarian  cell,  which  is  already,  as  a  rule,  in  process  of 
segmentation,  is  surrounded  by  numerous  large  yolk  cells ;  the 
proglottids  nearest  the  posterior  extremity  are  frequently  eggless. 

The   eggs,  which   are   deposited    in   the  intestine  and   evacuated 


^  Until  recently  this  worm,  which  was  understood  to  belong  to  a  separate  species, 
was  proved  on  examination  by  R.  Blanchard  ("Mai.  Par.,"  1896),  to  be  Dlbothrio- 
cephalus latus.  Compare  also  Galli-Valerio,  in  Centralbl.  f.  Bakt,,  Path,  und  In- 
fektionskr.,  1900  (i),  xxvii,  p.  308. 


DIBOTHRIOCEPHALUS   LATUS 


311 


with  the  faeces,  hatch  in  water  after  a  fortnight  or  more ;  the  embryonal 
integument  (embryophore)  of  the  oncosphere  is  provided  with  ciUa  ; 
after  bursting  open  the  Hd  of  the  egg  the  oncosphere  in  its  embryo- 
phore (fig.  207)  reaches  the  water  and  swims  slowly  about ;  often  it 
slips  out  of  its  ciliated  embryophore,  sinks  to  the  bottom  and  is  capable 
of  a  creeping  motion;  sooner  or  later  it  dies  in  the  water.  The 
manner  and  means  of  its  invasion  of  an  intermediate  host  are  still 


1 

r^ 

t 

a 

ril 

rf 

i?- 

tt 

m 

ffi 

33 

Fig.  204, — Various  chains  of 
segments  of  Dibothriocephalus 
la/us,  showing  the  central 
uterine    rosette.     (Natural   size.) 


Fig.  205. — Transverse  sec- 
tion  of  the    head    of  Dibo- 

thriocephahis  latus,     30/1. 


Fig.  206. — Fairly  mature  proglottis  of  Dibothriocephalus 
lat'iis.  The  vitellaria  are  at  the  sides  ;  the  uterus,  filled 
with  eggs,  is  in  the  middle,  also  the  vagina  (the  dark  stripe 
passing  almost  straight  from  the  front  to  the  back),  and  the 
vas  deferens  (almost  hidden  by  the  uterus).  Above  in  the 
centre  is  the  cirrus  sac,  and  below  the  shell  gland  and  ovary 
are  seen.     15/1.     (From  a  stained  preparation.) 


unknown ;  yet  we  are  aware  that  the  larval  stage  (plerocercoid, 
fig.  208),  which  resembles  the  scolex  and  may  reach  a  length  of  30  mm., 
lives  in  the  intestine,  in  the  intestinal  wall,  in  the  liver,  spleen,  genital 
glands  and  muscular  system  (fig.  209)  of  various  fresh-water  fish, 
the  pike  {Esox  lucius),  the  miller's  thumb  {Lota  vulgaris),  the  perch 
(Perca  fliwiatilis),  Salmo  umbla,  Triitta  vulgaris^  Tr.  lacustrls,  Thyntallis 
vulgaris  (grayling),  Coregonus  lavareUiSj  C.  albula  (in  Europe)  and 
Onchorhynchiis  perryi  (in  Japan).  The  transmission  of  the  plero- 
cercoids  from  these  fish  to  the  dog,  cat  and  man  (Braun,  Parona,  Grassi 
and  Ferrara,  Grassi  and  Rovelli,  Ijima,  Zschokke,  Schroeder)  leads  to 


,12 


THE   ANIMAL   PARASITES   OF   MAN 


the  development  of  the  broad  tapeworm,  the  growth  of  which  is  rapid. 
In  my  experiments  on  human  beings  the  average  number  of  pro- 
glottids  formed  per  diem  averaged  thirty-one  to  thirty-two  for  five 
weeks,  with  a  length  of  8  to  9  cm.     According  to  Parona  the  eggs 


Fig.  207. — Dibothriocephalus  lafus :  development  of  egg.  i,  segmentation  complete  ;  some 
cells  of  the  blastosphere  have  migrated  through  the  yolk  and  have  flattened  to  form  f,  the 
yolk  envelope  ;  others  form  a  layer  of  flattened  cells  (<;)  forming  the  embryophore  ;  the 
remaining  cells  {d)  of  the  blastosphere  form  the  hexacanth  embryo.  2,  embryophore  [e)  is 
becoming  thicker.  3,  the  ciliated  embryo  has  been  pressed  out  of  the  shell  ;  s\  the  operculum  ; 
^,  the  yolk  envelope  remaining  in  the  shell  [s) ',  j,  the  yolk  consisting  of  separate  cells.  4, 
a  free-swimming  larva  much  swollen  by  the  water.     (After  Benham  and  Schauinsland.) 


appear  twenty-four  days  after  man  has  been  infected.  Zschokke  found 
the  average  growth  in  the  experimental  infection  of  man  between  5*2 
and  8'2  cm.  per  diem,  and  the  person  experimented  upon  by  Ijima 
evacuated  a  piece  of  a  Dibothriocephalus  latus,  22*5  cm.  in  length, 
only  twenty-one  days  after  the  infection. 


DIBOTHRIOCEPHALUS   LATUS 


313 


The  "  broad  tapeworm "  is  a  frequent  parasite 
of  man  in  some  districts,  but  it  also  occurs  in  the 
domestic  dog,  and  on  rare  occasions  is  found  in 
the  domestic  cat  (together  with  Dibotliriocephahis 
felis^  Crepl.)  and  fox.  French  Switzerland  and  the 
Baltic  Provinces  of  Russia  are  the  centres  of  distri- 
bution ;  from  the  former  districts  the  distribution 
radiates  to  France  and  Italy  (Lombardy,  Piedmont) ; 
from  the  Baltic  Provinces  over  Ingermanland 
to  Petrograd,  over  Finland  to  Sweden  (on  the 
shore  of  the  Gulf  of  Bothnia),  in  a  southerly  direc- 
tion to  Poland,  and  into  the  Russian  Empire  and 
across  it  to  Roumania,  and  towards  the  west  along 
the  coast  of  the  Baltic  Sea  to  the  North  Sea,  where,  however,  its 
frequency  considerably  diminishes  (Holland,  Belgium,  and  the 
North  of  France). 

In  Turkestan  and  Japan  the  ^'  broad  tapeworm "  is  the  most 
frequent  parasite  of  man  ;  it  has  been  reported  in  Africa  from  the 
vicinity  of    Lake    N'gami   as  well    as    from    Madagascar  ;    cases,    in 


Fig.  208.— Plero- 
cercoid  of  Dibothrio- 
cephalus  latus.  A., 
with  the  head 
evaginated  ;  B.,  with 
the  head  invaginated. 
From  the  muscle  of 
the  pike. 


Fig.  209. — A  piece  of  the  body  wall  of  the  Burbot,  Lo/a  vulgaris.  The 
tangential  section  has  exposed  the  muscles  of  the  trunk,  with  a  plerocercoid  of 
Dibothriocephalus  latus.     Natural  size. 


part  at  least  imported,  have  also  come  under  observation  in  North 
America. 

In   Germany    Dihothriocephaliis   latus — apart    from   the   fact   that 
it   is   undoubtedly   imported   from   Switzerland,    Russia   or    Italy— is 


314  THE   ANIMAL   PARASITES   OF    MAN 

particularly  frequent  in  East  Prussia  amongst  the  inhabitants  of 
the  Courland  Lagoon  district,  on  the  Baltic  ;  it  is,  moreover,  also 
found  in  the  Province  and  even  in  the  City  of  Konigsberg.  In  West 
Prussia  and  Pomerania  it  is  very  much  scarcer. 

It  is  also  found  in  Munich  and  in  the  vicinity  of  the  Lake  of 
Starnberg  (Bollinger). 

Krabbe  found  it  in  10  per  cent,  of  the  sufferers  from  tape- 
worms in  Denmark;  Szydlowski  found  the  ova  of  this  worm  in 
Dorpat  in  10  per  cent,  of  the  faeces  examined;  Kruse  found  the 
worm  in  6  per  cent,  of  post-mortems  ;  Kessler,  in  Petrograd, 
found  the  eggs  in  the  faeces  in  7*8  per  cent. ;  at  post-mortems  he 
found  the  worms  in  1*17  per  cent.,  though  Winogradoff  only  found 
it  in  o'8  per  cent.  In  Moscow,  according  to  Baranovsky,  8*9 
per  cent,  of  the  faeces  examined  contained  the  ova  of  Dihothrio- 
cephalus.  In  the  interior  and  southern  provinces  of  Sweden  the 
worm,  according  to  Lonnberg,  is  only  found  sporadically,  but,  on 
the  other  hand,  in  Angermanland  about  to  per  cent,  of  the  popula- 
tion is  affected  ;  while  again  in  Norbotten  the  majority  of  persons 
are  affected,  and  in  Haparanda  the  entire  population  (with  the 
exception  of  infants)  harbour  this  parasite.  In  Switzerland 
D.  latiis  is  very  frequent  in  close  proximity  to  the  lakes  of  Bieler, 
Neuchatel,  Morat  and  Geneva  (according  to  Zaeslin  10  to  15  to 
20  per  cent,  of  the  population  are  affected)  ;  the  parasite  is  less 
frequent  in  districts  one  to  four  hours  removed  from  these  lakes. 

Of  the  fish  from  Swiss  lakes  examined  by  Schor  those  from 
Lake  Geneva  were  most  commonly  infected,  and  especially  Lota  sp. 
and  Perca  sp. 

The  frequency  and  distribution  have,  nevertheless,  decreased 
perceptibly  in  places;  at  the  commencement  of  the  eighteenth 
century  the  broad  tapeworm  was  very  common  in  Paris,  at  the 
present  date  it  only  occurs  when  imported  (Blanchard)  ;  in 
Geneva,  also,  according  to  Zschokke,  it  has  become  rarer  (formerly 
10  per  cent.,  now  only  i  per  cent.). 

The  disturbances  produced  in  man  by  the  presence  of  broad 
tapeworms  are,  as  a  rule,  very  trifling ;  in  other  cases  they  produce 
partly  gastric  disorders  and  partly  nervous  symptoms ;  in  a  number 
of  cases,  again,  they  set  up  severe  anaemia,  apparently  caused  by 
toxins  produced  by  the  worms  and  absorbed  by  the  host.  There 
is  no  danger  of  auto-infection,  as  the  larval  stage  lives  only  in 
fishes,  not  in  warm-blooded  animals.  The  case  reported  by 
Meschede  (ova  like  those  of  Dihothriocephalus  latus  in  the  brain  of 
a  man  who  had  suffered  from  epilepsy  for  six  years)  must  be  otherwise 
explained. 

Human    beings,  like    other    hosts,  can    only   acquire    the    broad 


DIBOTHRIOCEPHALUS   CORDATUS  3I5 

tapeworm  by  ingesting  its  plerocercoids  with  the  previously  men- 
tioned fresh-water  fishes  ;  the  opportunity  for  such  infection  is 
afforded  the  more  readily  by  the  fact  that  not  only  do  the  lower 
classes  not  pay  sufficient  attention  to  the  cooking  of  fish,  so  that 
all  the  larvae  that  are  present  may  be  killed,  but  also  in  certain 
localities  the  custom  exists  of  eating  some  parts  of  these  fishes  in  a 
raw  condition ;  even  the  mere  handHng  of  the  usually  severely 
infected  intermediary  hosts  may  occasionally  cause  infection.  The 
plerocercoids  are  as  well  known  as,  but  differ  materially  in  appear- 
ance from,  the  cysticerci  {Cysticercns  celhdosce)  of  pig's  flesh.  In 
Germany  the  occurrence  of  the  plerocercoids  of  Dihothriocephalus 
latus  has  been  confirmed  in  the  pike,  miller's  thumb  and  perch  of 
East  Prussia,  and  more  particularly  in  those  taken  from  the  Courland 
Lagoon. 

The  life  of  D.  latus  is  a  very  long  one  (six  to  fourteen  years), 
as  is  deduced  from  persons  who  have  left  D.  latus  regions  after  they 
have  been  infected. 

According  to  the  experiments  of  M.  Sciior,  plerocercoids  of 
D.  latus  placed  in  slowly  warmed  water  completely  lose  their  move- 
ment at  54*^  to  55'  C. ;  they  survive  the  death  of  their  host  for  several 
days  ;  they  are  killed  by  low  temperatures  -  3°  to  -h  1°  C.  in  two 
days  ;  strong  acids  and  salt  solutions  kill  them  at  once,  also  high 
temperatures,  but  all  the  same  at  least  ten  minutes  is  required  in 
boiling  or  frying  fish  in  order  to  kill  the  plerocercoids  with 
certainty. 


Dibothriocephalus  cordatus,  R.  Lkt., 
1863. 

Syn. :  Bothriocephalus  cordatus^  R.  Lkt. 

Length,  80  to  115  cm.  ;  the  head  is  heart-         Fig.  210.— Cephalic  end 
shaped    and    measures    2    by    2    mm.      The      °^  Dibothriocephalus  corda^ 

^  in  f  ^^^ '    °"    ^^^    '^"    Viewed 

suctorial  grooves  are  on  the  flat  surface  ;  the      sideways,  on  the  right  from 
segments  commence  close  behind  the  head  and      ^^^  dorsal  surface,  showing 

^  .  .  a  suctorial  groove.     (After 

increase  rapidly  in   breadth.     At   only  3    cm.      Leuckart.) 
behind  the  head  they  are  already  mature  ;  the 

greatest  breadth  attained  by  them  averages  7  to  8  mm.,  the  length  3  to 
4  mm. ;  the  number  of  proglottids  averages  600 ;  the  most  posterior  ones 
are  usually  square.  The  uterine  rosette  is  generally  formed  of  six  to 
eight  lateral  loops.  The  eggs  are  operculated  and  measure  75  /a  by  50  />t. 
Dibothriocephalus  cordatus  is  a  common  parasite  of  the  seal,  the 
walrus  and  the  dog  in  Greenland  and  Iceland,  occasionally  of  man 
also.     No  doubt  its  larva  lives  in  fishes. 


3l6  THE   ANIMAL   PARASITES   OF   MAN 

The  statement  that  D.  cordatus  also  occurs  in  Dorpat  in  human  beings 
has  been  proved  erroneous  {Zool.  Anzeiger^  1882,  v,  p.  46),  as  also  has  the 
report  that  this  worm  lives  in  hares  in  the  neighbourhood  of  Berlin,  whither  it 
was  supposed  to  have  been  carried  by  Esquimaux  dogs  (Rosenkranz  in  Deutsch. 
med.  Wochenschr.^  ^^77,  iiij  P*  620).  The  parasite  stated  by  the  author  to  i)e 
D.  cordatus  is  Tcenia  pectinata,  Goeze,  which  has  been  known  since  1766. 

Dibothriocephalus  parvus,  Stephens,  1908. 

Largest  gravid  segments  5  by  3  mm.  Uterus  forms  a  central 
rosette  with  four  to  five  loops  on  each  side  of  median  Hne.  In  a  pro- 
glottid measuring  3*5  by  2*25  mm.  the  genital  atrium  is  situated  0*4 
to  o'5  mm.  behind  the  anterior  margin  and  the  uterine  opening  the 
same  distance  behind  the  genital  atrium.  Calcareous  corpuscles  absent 
in  the  preserved  specimens.     Eggs  operculated,  59*2  /x  by  407  /x. 

Distinguished  from  Dibothriocephalus  latus — (i)  by  the  size  of  gravid 
segments  (the  minimum  width  of  gravid  segments  of  D.  laUis  is  10 
to  12  mm.,  so  that  D.  parvus  is  a  much  smaller  worm)  ;  (2)  quadrate 
segments  of  D.  latus  measure  6  by  6  mm.,  those  of  D.  parvus  4  by 
4  mm.  ;   (3)  by  the  eggs. 

From  D.  cordatus  it  is  distinguished  by — (i)  D.  cordatus  has  only 
fifty  immature  segments,  D.  parvus  has  at  least  200,  possibly  more  ; 
(2)  mature  segments  of  D.  cordatus  measure  7  to  8  mm.,  maximum 
width  of  D.  parvus  is  5  mm. ;  (3)  quadrate  segments  of  D.  cordatus 
measure  5  to  6  mm. ;  (4)  D.  cordatus  has  six  to  eight  uterine  loops ; 
(5)  Z).  cordatus  measures  75/1,  to  80//,  by  50//.. 

Habitat. — Intestine  of  man  (Syrian,  in  Tasmania). 

Genus.     DIplogonoporus,   Lonnbrg.,  1892. 
Syn.:  Krabbea^  R.  Blanch.,  1894. 

The  scolex  is  short  and  has  powerful  suctorial  grooves  ;  no  neck  ; 
the  proglottids  are  short  and  broad ;  there  are  two  sets  of  genital 
organs  side  by  side  in  each  segment,  which  in  all  essentials  resemble 
the  single  one  of  Dibothriocephalus. 

Parasitic  in  whales  and  seals,  occasionally  in  man. 

Diplogonoporus  grandls,  R.  Blanch.,  1894. 

Syn. :  Bothriocephalus  sp.,  Ijima  et  Kurimoto,  1894  ;  Krabbea  gra?tdis,  R.  Blanch. 

Scolex  unknown  ;  chain  of  proglottids  over  10  m.  in  length, 
1*5  mm.  broad  anteriorly,  25  mm.  broad  posteriorly.  The  proglottids 
are  very  short  (0*45  mm.),  but  14  to  16  mm.  broad.  On  either  side  to 
the  right  and  left  of  the  worm,  along  the  entire  ventral  surface,  there 
is  a  longitudinal  groove  ;  these  grooves  are  nearer  to  each  other  than 


SPARGANUM 


317 


to  the  lateral  margin  ;  in  them,  lie  the  genital  pores,  and  they  are  in 
the  same  sequence  as  in  Dibothriocephalns ;  corresponding  to  the 
scanty  length  (0-45  mm.)  of  the  proglottids,  the  ovary  is  only  developed 
transversely  ;  the  uterus  only  makes  a  few  loops.  Eggs  (fig.  195) 
thick  shelled,  brown,  63  yu,  by  48  yit  to  50  fju.  This  parasite  has  hitherto 
been  observed  twice  in  Japanese.  Similar  species  are  known  in  Cetacea 
and  seals. 

; 


Fig.  211. — Diplogonoporus  grandis,  Llihe,  1899:  ventral  view  of  a  portion  of  the 
strobila,  showing  two  rows  of  genital  pores  and  partially  extruded  cirri.  (After  Ijima  and 
Kurimoto.) 


^Z€t-^ 


zct. 


^==^5^ 


Fig.  212. — Diplogonoporus  grandis  :  ventral  view  (diagrammatic)  of  genitalia 
of  left  side  ;  cir,  cirrus  ;  cir.o,  cirrus  opening  ;  dtg.^  vitelline  duct ;  ov.^  ovary  ; 
ovd.,  oviduct ;  j^.,  receptaculum  seminis ;  w/.,  uterus;  ut.o.,  uterine  pore  ;  vag.y 
vagina;  vag.o.,  vaginal  pore;  vd,  vas  deferens.  X  150.  (After  Ijima  and 
Kurimoto.) 


Sparganum,  Diesing,  1854. 

The  term  S^ar^^;///m,  invented  by  Diesing,  is  used  as  a  group  name 
of  larval  bothriocephalid  Cestodes  whose  development  is  not  suffi- 
ciently advanced  to  enable  them  to  be  assigned  to  any  particular  genus. 

Sparganum  mansoni,  Cobb.,   1883. 

Syn. :  Ligula  mansoni,  Cobbold,  1883;  Bothriocephalus  linguloides,K.  Lkt., 
1886  ;  Bothriocephalus  ?nansoni,  R.  Blanch.,  1886. 

These  plerocercoids  were  discovered  in  1882  by  P.  Manson 
during   the  post-iuoriem  on    a    Chinaman   who    had   died  in  Amoy, 


3i8 


THE   ANIMAL   PARASITES   OF   MAN 


Fig.  213.  —  Ce- 
phalic end  of  Sparga- 
num  mansoni,  Cobb. 
{After  Leuckart.) 


twelve  Specimens  being  found  beneath  the  peri- 
toneum and  one  free  in  the  abdominal  cavity. 
Cobbold  described  them  as  Ligula  uiaiisoni,  and 
Leuckart,  who  contemporaneously  reported  a  case 
in  Japan,  termed  them  BothriocephaUis  liguloldes. 
Ijima  and  Murata  reported  eight  further  cases,  and 
Miyake  records  nine  further  cases,  seven  of  which 
are  recorded  in  Japanese  literature. 

The  plerocercoid,  which  hitherto  alone  is  known 
to  us,  attains  a  length  of  30  cm.  and  a  breadth  of 
3  to  6  to  12  mm.  The  ribbon-shaped  body  is 
wrinkled,  the  lateral  borders  are  often  somewhat 
thickened,  so  that  the  transverse  section  has  the 
form  of  a  biscuit ;  the  anterior  end  is  usually  w^ider 
and  has  the  head  provided  w^ith  two  weak  suctorial 
grooves,  either  retracted  or  protracted. 

The  parasite  makes  migrations  within  the  body, 
and  thus  may  reach  the  urinary  passages ;  then  it  is 
either  evacuated  with  the  urine  or  has  to  be  removed 
from  the  urethra ;  not  rarely  it  causes  non-infiam- 
matory  tumours  on  various  parts  of  the  skin,  which 
are  at  times  painful  and  at  times  vary  in  size. 

Nothing  is  knov/n  of  its  development  and  origin. 


Sparganum  proliferum,  Ijima,   1905. 

Syn. :  Plerocercoides prolifer,  Ijima,  1905  ;  Sparganum 
prolifer^  Verdun,  Man  son,  1907. 

These  plerocercoids  produce  an  acne-like  con- 
dition of  the  skin.  The  condition  is  really  one  of 
capsules  in  great  abundance  in  the  subcutaneous 
tissue  and  less  so  in  the  corium  and  in  the  inter- 
muscular connective  tissue.  The  encapsuled  worms 
in  the  corium  feel  like  embedded  rice  grains  and 
raise  the  epidermis,  giving  rise  to  an  acne-like  con- 
dition. Many  thousands  occur  scattered  over  the 
body;  in  Ijima's  Japanese  case  there  were  over 
10,000  in  the  left  thigh.  The  worms  when  they  first 
appear  in  the  skin  cause  itching.  The  capsules  are 
ovoid,  generally  about  i  to  2  mm.  in  diameter,  but 
they  may  be  smaller  and  also  much  larger.  The 
larger  ones  occur  in  the  subcutaneous  tissue.  The 
capsules  consist  of  dense  tough  connective  tissue. 
Each  capsule,  as  a  rule,  contains  one  worm,  but  as  many  as  seven 
may  occur.     The  skin  of  areas  that  have  been  long  infected  is  swollen 


Fig.  21^.— Sparga- 
num mansoni :  on 
the  right  in  transverse 
section.  Natural  size. 
(After  Ijima  and 
Murata.) 


SPARGANUM    PROLIFERUM 


3^9 


V 


A  A 


and  indurated  or  adherent,  giving  a  somewhat  elephantoid  appearance. 
The  subcutaneous  tissue  is  thick  and  filled  with  slimy  fluid  or  in 
other  parts  sclerosed. 

The  Worm. — The  chief  peculiarity  is  its 
irregular  shape  and  its  reproduction  in  the 
larval  stage  by  forming  supernumerary  heads, 
which  are  supposed  to  wander  about  the 
body. 

The  simplest  forms  are  thread-like  bodies, 
flat  or  round,  3  mm.  long  and  0*3  mm.  in 
diameter,  but  they  may  be  12  mm.  long  by 
2'5  mm.  broad.  The  narrow  end  is  the  head, 
which  in  life  invaginates  and  evaginates,  but 
there  is  no  indication  of  any  suckers,  except 
an  inconstant  terminal  depression.  In  addi- 
tion to  these  simple  forms  the  most  compli- 
cated and  irregular  forms  occur,  due  to  the         ^^^-  ^is.—Sparganumproii- 

c  \-  r    1      J       /u       J   X       i.  •  L  f'^  '  •*^"  ^'^^    t)uds,  right   ex- 

tormation  of  buds  (heads)   at  various  parts,      tended,     x  4.    (After  ijima.) 


"^,1 


Fig.  216. — Sparganmn ptolifeiii7n,      x   lo.     (After  Stiles.) 

The  detachment  and  growth  of  a  head  account  for  the  presence  of 
more  than    one   worm  in  a  cyst.     The  irregularity  in    form  is   also 


320 


THE   ANIMAL   PARASITES   OF   MAN 


increased  by  the  presence  in  the  subcuticular  tissue  of  the  worm  of 
reserve  food  bodies.  These  bodies  are  supposed  to  be  of  this  nature 
and  are  spherical,  lOo  /x  to  300  fju  in  diameter,  but  also  much  elongated. 

Calcareous  bodies  in  the  Japanese  worms  were  7*5  /-t  to  12  /^ ;  in  the 
Florida  worms  8-8  fi  to  17*6  jx. 

Mode  of  Infection. — Probably  from  eating  uncooked  fish. 

Distribution. — Japan,  Florida. 

Family.     Dipylidiidae,  Liihe,  1910. 

Genus.     DIpylidium,  R.  Lkt.,  1863. 

Rostellum  retractile,  with  several  rings  of  alternating  hooks  ;  the  latter  with  a 
disc-like  base,  having  the  shape  of  the  thorns  of  a  rose.  Genital  pores  opposite ; 
genitalia  double.  Testes  very  numerous  in  the  central  field ;  ovary  with  two  lobes  ; 
the  vitellaria,  which  are  smaller,  behind  them  ;  the  uterus  forms  a  reticulum,  in  the 
network  of  which  the  testicular  vesicles  lie  ;  later  on  it  breaks  up  into  sacs  enclosing 
one  or  several  eggs.     The  eggs  have  a  double  shell. 


\ 


DIpylidium  caninum,  L.,  1758. 

Syn.  :  Tcenia  ca?thta,  L.,  1758,  p.  p.  ;  Tcznia 
moniliformis^  Pallas,  1781;  Tcenia  ciicu7nerina, 
Bloch,  1782;  Tcenia  elliptica,  Batsch,  1786; 
DIpylidium  cucimierimim^  Lkt.,  1863. 

This  worm  measures  15  to  35  cm. 
in  length  and  1*5  to  3  mm.  in  breadth. 
The  scolex  is  small,  rhomboidal,  and 
has  a  club-shaped  rostellum  on  which 


hzz:i 


Fig.  217. — Dipylidium  caninum :  on  the 
left,  the  scolex,  neck  and  the  first  proglot- 
tids ;  on  the  right,  at  the  top,  a  packet  of 
ova  ;  below,  hooks  of  the  rostellum,  side 
and  front  views ;  below,  an  ovum.  Various 
magnifications.     (After  Diamare.) 


Fig.  218. — Dipylidium  caninum  '.  egg  show- 
ing a,  egg-shell  (vitelline  membrane  of  Moniez)  j 
b,  albuminous  coat ;  c,  internal  shell  formed  of 
or  secreted  by  an  outer  layer  of  blastomeres 
(Moniez)  ;  d^  hexacanth  embryo.  (After  Benham 
and  Moniez.) 


there  are,  in  three  to  four  rings,  forty-eight  to  sixty  hooks  resembling 
rose  thorns,  the  size  of  those  in  the  foremost  being  1 1  />t  to  15  /^  and  those 
in  the  hindmost  ring  6  //,.  The  neck  is  very  short,  the  most  anterior 
segments  broad  and  short,  the  middle  as  long  as  they  are  broad  ;  the 


DIPYLIDIUM   CANINUM 


321 


Fig.  219. — Dipylidium  caninum  :  central  portion  of  a  proglottis.  C.p.,  cirrus 
sac;  V.S.,  vitellaria  ;  ^Jtr.z'.,  excretory  vessels  ;  7".,  testicles  lying  in  the  meshes 
of  the  uterine  reticulum  which  laterally  forms  pouches  ;  O.,  ovary  ;  C/.,  reticulum 
of  uterus  ;  V.,  vagina  and  seminal  receptacle  (below  ovary).  Magnified.  '_  (After 
Neumann  and  Kailliet.) 


Fig.  220. — Dipylidium  caninum  :  development  of  embryo,  i,  solid  hexacanth  embryo  ; 
2,  primitive  lacuna  {a)  in  the  embryo  ;  3,  elongation  of  hinder  part,  rudiments  of  sucker  and 
rostellum  appearing ;  4,  "  body  "  and  "  tail "  distinct,  (d)  and  (c)  excretory  system  ;  5,  fore-body 
invaginates  into  hind-body,  excretory  bladder  has  a  pore;  6,  tail  has  dropped  off;  scolex 
growing  up  into  secondary  cavity  formed  by  fore-body  ;  the  primitive  cavity  has  been  absorbed 
at  stage  4.     (After  Benham,  Grassi  and  Rovelli.) 

21 


322 


THE   ANIMAL   PARASITES    OF   MAN 


/ 


mature  segments  are  longer  than  wide  (6  to  7  mm.  by  2  to  3  mm.),  fairly 
thick,  are  frequently  of  a  reddish  colour,  and  when  cast  off  resemble 
cucumber  seeds.  The  genital  pores  lie  symmetrically  at  the  lateral 
margins ;  the  roundish  egg  sacs,  arising  from  the  uterine  reticulum, 
contain  eight  to  fifteen  eggs  embedded  in  a  reddish  cement  substance 

(in  life).     The  eggs  are  globu- 

/.^'"vr— ^— ->>.  lar    (43   /t  to   50  /t)  ;  the  em- 

^^  bryonal  shell  (embryophore)  is 

^^^ ^  thin,  the  oncosphere  measures 

32 /i  to  36/1,.  Surrounding  the 
embryophore  is  an  albumin- 
ous coating,  and  outside  this 
the  thin  vitelline  envelope 
(fig.  218). 

Dipyiidiuin  caninuni  is  a 
common  intestinal  parasite  of 
dogs,  in  which  it  grows  larger 
{Tccnia  ciicnmerlna,  Bloch) 
than  in  cats  {T.  elliptical 
Batsch) ;  it  has,  however,  also 
been  found  in  jackals,  as  well 
as  in  human  beings,  though 
in  the  latter  it  is  of  compara- 
tively rare  occurrence  (twenty- 
four  cases),  and  almost  always 
affects  children,  generally  of 
tender  age.  One-third  of  all 
the  cases  in  children  were 
sucklings,  about  a  quarter  of 
all  the  cases  recorded  were 
adults,  and  these  occurred 
throughout  all  Europe  with 
the  exception  of  Spain  and 
Italy. 

The  proglottids,  which 
leave  the  intestine  spontane- 
ously, are  recognizable  by  the 
naked  eye  on  account  of  their 
form  and  reddish  colour,  as 
As  a  rule,  the  presence  of  this  parasite 


Fig.  221. — Larva  (cyslicercoid)  of  Dipylidium 
caninum^  consisting  of  body  and  tail.  The  latter 
is  solid  and  bears  on  it  the  embryonal  spines.  The 
bladder,  which  was  only  slightly  developed,  has 
disappeared,  and  the  fore-part  of  the  body  bearing 
the  rostellum  is  now  seen  invaginated  into  the 
hind  portion.  The  booklets  are  shown  in  front 
of  the  excretory  system  which  has  now  developed. 
At  a  further  stage  the  tail  drops  off;  the  head  now 
evaginates,  but  is  still  enclosed  in  a  double-walled 
sac  formed  by  the  prolongation  upwards  on  each 
side  of  the  topmost  parts  of  the  body  shown  in 
the  figure.  Cf.  fig.  220,  6.  Enlarged.  (After 
Grassi  and  Rovelli.) 


well  as  their  two  genital  pores. 

sets  up  no  marked  symptom  in  the  patient. 

The  corresponding  larval  form  (cysticercoid)  lives  in  the  louse 
of  the  dog  {Tricho elect es  canis),  a  fact  that  was  first  established  by 
Melnikow   and    Leuckart ;    according  to  Grassi  and  Rovelli,  as  well 


HYMENOLEPIDID^  323 

as  Sonsino,  it  also  lives  in  the  flea  of  the  dog  (Ctenocephalus  cants) 
and  in  the  flea  of  man  (Pulex  irritans),  but  not  in  its  larva.  The 
adult  segments,  which  also  leave  the  rectum  of  dogs  and  cats 
spontaneously,  creep  about  around  the  anus  and  get  into  the  hair, 
and  are  thus  partly  dried  and  disintegrated.  Part  of  the  segments,  or 
the  oncospheres  released  by  disintegration,  are  then  taken  up  by  lice 
and  fleas,  within  which  they  develop  into  larvae  (cysticercoids).  Dogs 
and  cats  are  thus  infected  by  then-  own  skin  parasites,  which  they  bite 
and  swallow  whilst  gnawing  at  their  fur.  The  infection  of  human 
beings  must  occur  in  an  analogous  manner,  by  transmission  of  the 
cysticercoids  present  on  the  lips  or  tongue  of  dogs  when  the  latter  lick 
them,  or  it  may  be  that  the  vermin  of  cats  and  dogs  harbouring  cysti- 
cercoids are  accidentally  and  directly  swallowed  by  human  beings. 


Family.     Hymenolepididae,  Railliet  and  Henry,  1909. 
Genus.     Hymenolepis,^  Weinland,  1858. 

Accessory  sac  (opening  into  genital  atrium)  usually  absent.  Vas  deferens  with 
an  external  (outside  cirrus  sac)  and  an  internal  (inside  cirrus  sac)  "seminal  vesicle." 
Three  testes  in  each  proglottis.  The  eggs  are  round  or  oval  with  two  to  four  distinct 
envelopes.     In  mammals  and  birds. 

Hynnenolepis  nana,  v.  Sieb.,  1852. 

Syn.  :  Tcenia  nana,  v.  Sieb.,  1852,  nfc  van  Beneden,  1867;  Tcenia  cegyptiaca, 
Bil.,  1852  ;  Diplacanthus-nanus,  VVeinld.,  1858  ;  Tcettia  {Hy?nenolepts)  nana,  Lkt., 
1863. 

The  worm  is  10  to  45  mm.  in  length  and  0*5  to  07  mm.  in  breadth ; 
the  head  is  globular,  0*25  to  0*30  mm.  in  diameter.  The  rostellum 
has  a  single  circlet  consisting  of  twenty-four  or  twenty-eight  to  thirty 
hooks,  which  are  only  14  /x  to  18  yu,  in  length.  The  neck  is  moderately 
long;  the  proglottids  are  very  narrow,  up  to  200  in  number, 
0*4  to  o*9  mm.  in  breadth,  and  0*014  *o  0*030  mm.  in  length.  The 
eggs  are  globular  or  oval,  30  /^  to  37  /a  to  48  /a  ;  the  oncospheres  measure 
16  yLt  to  19  /x  in  diameter,  with  two  coats,  separated  by  an  intervening 
semi-fluid  substance  (fig.  224). 

This  species  was  discovered  by  Bilharz  in  Cairo  in  185 1  ;  it  was 
found  by  him  in  great  numbers  in  the  intestine  of  a  boy  who  had 

'  The  genus  is  by  some  authors  divided  into  two  sub-genera— Hymenolepis,  s.  str.,  and 
Drepanidotaenia,  Raill. 

Drepanidotrsnia.— Body,  broad  lanceolate,  testes  three,  female  genitalia  antiporal  beside 
the  testes.  Scolex  small,  with  eight  hooks.  Neck  very  short,  longitudinal  muscle  bundles 
very  numerous.     No  accessory  sac  opening  into  genital  atrium. 

Hymenolepis.— ^d^xxovf ,  female  genitalia  ventral  to  or  between  testes. 


324 


THE   ANIMAL   PARASITES   OF   MAN 


died  of  meningitis.  For  several  years  this  was  the  only  case,  until 
1885,  since  when  numerous  cases  have  come  to  light.  Spooner  (1873) 
even  reported  a  case  from  North  America,  which  may,  however,  have 
related  to  Hynienolepis  diminuta.  In  Europe  the  w^orm  is  particu- 
larly frequent  in  Sicily,  but  it  has  also  been  repeatedly  observed  in 
North  Italy;  it  has,  moreover,  been  reported  from  Russia,  Servia, 
England,  France,  Germany,  North  and  South  America,  the  Philippines, 


Fig.  223.  —  Hymenolepis 
nana:  head.  Enlarged. 
(After  Mettens.) 


Fig.  222. —  Hy- 
menolepis nana,  v. 
Sieb.  About  12  i. 
(After  Leuckart.) 


Fig.  224. — Hymenolepis 
nana  :  an  egg.  Highly  mag- 
nified.    (After  Grassi.) 


Fig.  225.  —  Longitudinal 
section  through  the  intes- 
tinal villus  of  a  rat,  with  the 
larva  (cysticercoid)  of  Hy- 
meno/epis  mtirina.  Magni-. 
fied.  (After  Grassi  and 
Rovelli.) 


Siam  and  Japan,  in  all  over  100  cases.  Notwithstanding  its 
small  size  this  worm  causes  considerable  disorders  in  its  hosts — 
mostly  children — as  it  sets  up  loss  of  appetite,  diarrhoea,  various 
nervous  disturbances,  and  even  epilepsy ;  all  these  symptoms,  how- 
ever, disappear  after  the  expulsion  of  the  parasites,  w^hich  are  generally 
present  in  large  numbers. 

The  development  as  well  as  the  manner  of  infection  is  still 
unknown ;  Grassi  is  of  opinion  that  Hymenolepis  nana  is  indeed 
merely  a  variety  of  Hymenolepis  mnrina,  Duj.,    which   lives  in  rats. 


HYMENOLEPIS   NANA 


325 


According  to  Grassi  direct  development  takes  place  with  omission 
of  the  intermediate  host,  but  with  the  retention  of  the  larval  stage  ; 
that  is  to  say,  rats  infect  themselves  directly  with  Hymenolepis  iniirlna, 
by  ingesting  the  mature  segments  or  oncospheres  of  this  species,  from 
which  subsequently  the  small  larvae   originate  in  the  intestinal  wall 


cqrtpar: 


ry?'/.n., 


ex.Cdn. 


:M: 


ov. 


Fig.  226.  —  Hymenolepis  nana  (murma) :  cross  section  of  proglottis  from  a  rat.  c.p., 
cirrus  sac;  rec.  sem.,  receptaculum  seminis ;  s.^.,  shell  gland;  ov.,  ovary  ;/.,  testis  ;  corL 
par.,  cortical  parenchyma;  m.l.n.,  main  lateral  nerve;  ex.  can.,  excretory  canal;  y.^., 
vitellarium.     (After  v.  Linstow.) 


sec.cay^ 


■'pr.cay. 


pr.ca\/. 


':Caud. 


¥iG.22'j.— Hymenolepis  nana:  longitudinal  section  of  an  embryo,  bl.p., 
anterior  opening  of  secondary  cavity  ;  caud.,  caudal  appendage  ;  pr.  cav.,  primary 
cavity  ;  sec.  cav.,  secondary  cavity.     Enlarged.     (After  Grassi  and  Rovelli.) 

(fig.  225);  when  fully  developed  they  fall  into  the  intestinal  lumen 
and  become  tapeworms.  The  identity  of  the  two  forms  has  neverthe- 
less been  disputed  (Moniez,  R.  Blanchard,  v.  Linstow),  though  their 
near  relationship  cannot  be  denied.  Grassi  gave  mature  segments  of 
Hymenolepis  nturina  to  six  persons,  but  only  one  person  evacuated 
a  tapeworm.       This,  however,  proves    nothing   in    a    district  where 


326 


THE   ANIMAL   PARASITES   OF   MAN 


Hymenolepis  nana  frequently  occurs  in  man  ;  it  was,  moreover,  not 
possible  to  infect  rats  with  segments  of  Hymenolepis  nana  (of  man). 
Accordingly  this  form  may  represent  an  independent  species,  which, 
however,  like  Hymenolepis  murina,  also  omits  an  intermediate  host. 


Hymenolepis  diminuta,  Rud.,  1819. 

Syn.  :  TcEuia  diminuta^  Rud.,  1819  ;  Tcenia  leptocephala^  Crepl.,  1825  ;  TcEnia 
flavopU7ictata,  Weinld.,  1858  ;  Tcenia  varesina,  E.  Parona,  1884  ;  Tcenia  7niftima, 
Grass!,  1886. 

This  species  measures  20  to  60  cm.  in  length,  and  up  to  3*5  mm. 

in  breadth  ;  there  are  from  600  to  1,000  segments.     The  head  is  very 

small  (0*2  to  o*5  mm.),  it  is  club-shaped  and    has 

"'""^^^^^^  a  rudimentary  unarmed  rostellum;  the  neck  is  short ; 

^^  the  mature  segments  are  3*5  mm.  in  breadth,  066  mm. 

{  >  I        in  length  ;  the  eggs  are  round  or  oval.     The  outer 

^  egg-shell  is  yellowish  and  thickened,  with  indistinct 

'^  radial  stripes  ;  the  inner  embryonal  shell  (embryo- 

\  phore)   double,   thin  ;    the  outer  layer  is  somewhat 

pointed   at  the  poles ;    oncosphere   28  /x  by  36  fju. 

Between   the    inner  and    outer  shells   is  a   middle 

granular  layer. 

Hymenolepis  diminuta  lives  in  the  intestine  of 
rats — Mus  dectimanus  (the  sewer  rat),  Mus  ratins  (the 
black  rat),  and  Mus  alexandrimis,  rarely  in  mice  ;  it 
is  occasionally  also  found  in  human  beings. 

Weinland  described  it  from  specimens  collected 
by  Dr.  E.  Palmer  in  1842,  in  Boston,  from  a  child 
aged  19  months,  as  T.  flavopunctata.  A  second  case 
relating  to  a  three  year  old  child,  from  Philadelphia, 
was  only  reported  in  1889  by  Leidy ;  a  third  case 
was  simultaneously  reported  of  a  two  year  old  girl 
in  Varese  (T.  varesina)  ;  and  Grassi 
described  another  case  relating  to  a 
twelve  year  old  girl  from  Catania 
(Sicily).  Sonsino  and  Previtera  reported 


Fig.  228.  —  Hy- 
menolepis diminuta  ; 
scolex.  Magnified. 
(After  Zschokke.) 


P^IG.  229. — Hymenolepis  diminuta  :  two  pro 
glottids  showing  testes  (3),  ovary  and  vagina 
Slightly  enlarged.     (After  Grassi.) 


Fig.  230. — Hymenolepis   diminuld. 
egg  from  man.     (After  Bizzozero.) 


HYMENOLEPIS   DIMINUTA 


327 


the  same  species  in  Italy,  Zschokke  in  France,  Lutz  and  Magalhaes 
in  South  America,  and  Packard  in  North  America  :  a  total  of  twelve 
cases,  five  from  America,  the  rest  from  Europe  (Ransom). 

According  to  Grassi  and  Rovelli  the  larval  stage  lives  in  a  small 
moth  (Asopia  farinaiis),  as  well  as  in  its  larva,  in  an'  orthopteron 
{Anisolabis  anmilipes),  and  in  coleoptera  {Acis  spinosa  and  Scaurus 
striatus).     Experimental  infections  have  been    successful  on  rats  as 


/,  — -^ 


Fig.  231. — Hymenolepis  diminuta:  cyslicercoid  from  the  rat  ^Qdi{Ceraiophyllus  fasciatus) 
a,  remains  of  primary  vesicle  ;  d,  fibrous  layer  ;  c,  radially  striated  layer  resembling  cuticle  ; 
d,  layer  of  columnar  cells  ;  e,  parenchymatous  layer  of  irregularly  disposed  cells  ;  f,  parenchy- 
matous layer.     (Stephens,  after  Nicjll  and  Minchin.) 

well  as  on  human  beings.     In  America  other  species  of  insects  may 
be  the  intermediary  hosts. 

Nicoll  and  Minchin^  found  in  the  body  cavity  of  4  per  cent, 
of  rat  fleas  {Ceratophyllus  fasciatus)  the  cysticercoid  o  ^Hymenolepis 
diminuta.  That  it  belonged  to  this  species  was  shown  by  its  unarmed 
rostellum  and  by  feeding;  340  fleas  were  fed  to  white  rats  and  fourteen 


'  Proc.  Zool.  Soc,  191 1,  p.  9. 


328 


THE   ANIMAL   PARASITES   OF   MAN 


worms  obtained,  i.e.,  about  4  per  cent.,  thus  corresponding  to  the  infec- 
tion of  the  fleas.  The  development  in  the  flea  probably  begins  in  the 
pupal  stage,  the  eggs  being  ingested  by  the  older  flea  larvae.  The  larva 
is  o'3i  by  0*25  mm.;  tail  o'8  mm.,  scolex  0*075  by  0*09  mm.,  suckers, 
0*055  mm.  in  diameter.  Microscopically  it  shows — (i)  externally 
a  radially  striated  layer  resembling  cuticle,  (2)  a  layer  of  columnar  cells, 
(3)  parenchymatous  layer  continuous  with  the  tail,  (4)  fibrous  layer 
around  the  small  caudal  vesicle,  then  the  parenchymatous  scolex  at 
the  bottom  of  the  secondary  cavity. 

Nicoll  and  Minchin  (loc.  cit.)  found  a  cysticercoid  ^  in  the  rat  flea 
Ceratophyllus  fasciatus  which  w^as  probably  that  oi  Hyiuenolepis  miirina. 
Body  o-i6  mm.,  tail  o'lg  mm.,  scolex  0-096  mm.  in  diameter.  Ros- 
tellum  has  twenty-three  spines  in  a  single  row.  Length  0*017  mm., 
handle  o-oi  mm., guard  0*007  i^tn.,  prong  0*007  mm.  Sucker  0*042  mm. 
Although  this  cycle,  then,  for  H.  miirina  also  exists,  it  is  not  probable 
that  rats  (or  man  in  the  case  of  H.  nana  if  this  be  considered  distinct) 
infect  themselves  in  this  way,  as  they  hardly 
ingest  all  the  necessary  fleas  to  account  for  the 
massive  infection  which  frequently  exists  in 
rats  (and  man),  so  that  Grassi's  cycle  holds 
good  as  the  predominant  method.  Xenopsylla 
cheopis  has  also  been  found  by  Johnston  to 
harbour  both  cysticercoids  in  Australia. 

Hymenolepis  lanceolata,  Bloch,  1782. 

Syn.  :    To'm'a  lanceolata,  Bloch,  1782;   DrepanidotcEnia 

lanceolata,  Railliet,  1892. 

The   parasite  measures  30  to  130  mm.  in 

^~;      length    and    5   to    18    mm.    in    breadth;    the 

head      is     globular     and     very     small ;      the 


rec.  33/77. 


Fig.  232.  —  Hymenolepis 
lanceolata.  Natural  size. 
(After  Goeze.)  To  the 
right  above,  two  hooks. 
120/1.     (After  Krabbe.) 


Fig.  233. — Hymenolepis  lanceolata :  diagram  of 
female  genitalia,  ov.,  ovary;  ovd.,  oviduct;  rec.  sem., 
receptaculum  seminis  ;  s.g.,  shell  gland;  ut.,  uterus; 
y.g.,  vitellarium.     (After  Wolflhiigel.) 


'A  third  cysticercoid  resembling  this,  but  without  hooks,  has  also  been  found. 


DAVAINEID^ 


329 


rostellum  is  cylindrical,  with  a  circlet  composed  of  eight  hooks  (31  /x 
to  35  yit  in  length).  The  neck  is  very  short.  The  short  segments 
increase  gradually  and  equally  in  breadth,  but  only  a  little  in 
length  ;  the  female  glands  lie  on  the  side  opposite  to  that  on  which 
the  genital  pore  is  situated;  the  three  elliptical  testes  are  on  the 
same  side  as  the  pores  ;  the  cirrus  is  armed  and  slender.  The 
eggs  have  three  envelopes  and  are  oval  (50  /^  by  35  ytt),  the  external 
envelope  is  thin,  the  middle  intermediate  layer  or  envelope  is  not 
so  marked  as  in  H.  dijjiiniita,  and  the  internal  one  is  very  thin  and 
sometimes  has  polar  papillae,  as  in  Hymenolepis  diminuta  and 
H.  nana. 

It  inhabits  the  intestine  of  the  following  birds  :  Domesticated 
ducks  and  geese,  the  Muscovy  duck  (Cairlna  moschata),  white-headed 
duck  {Erisinatura  leticocephala),  the  pochard  (Nyroca  nifina),  and 
the  flamingo  (Phcenicopteriis  antiqnonim).  It  has  been  recorded  from 
Great  Britain,  France,  Denmark,  Austria  and  Germany. 

Zschokke  reports  the  receipt  of  two  specimens  which  a  twelve 
year  old  boy  in  Breslau  evacuated  spontaneously  at  two  different 
ti  mes. 

The  corresponding  larva,  according  to  Mrazek,  lives  in  fresh  water 
Cyclops;  according  to  Dadai  it  is  likewise  found  in  another  copepod, 
Diaptomus  spinosus,  but  the  hooks  of  Dadai's  larva  differed  in  size. 


Family.     Davaineidae,  Fuhrmann,  1907. 

Sub-family.     Davaineinae,  Braun,  1900. 

Genus.   Davainea,  R.  Blanch.,  1891. 

The  large  scolex  is  more  or  less  globular,  much  wider  than  the  rostellum,  which  is 
furnished  with  two  rings  of  very  small  and  numerous  hooks.  Neck  absent,  pro- 
gloltids  few,  genitalia  single.     Parasitic  chiefly  in  birds.' 

Davainea  madagascariensis,  Davaine,  1869. 

Syn. :   T<^nia  madagascariensis^  Dav. ;   Tcznia  demerariensis^  Daniels,  1895. 

This  worm  measures  25  to  30  cm.  in  length;  the  head  has  four 
large  round  suckers ;  the  rostellum  has  ninety  hooks  (18  /m  in  length)  ; 
there  are  500  to  700  segments,  of  which  the  last  100  are  filled 
with  eggs  and  form  half  of  the  entire  worm.  The  segments,  when 
mature,  measure  2  mm.  in  length  by  1*4  mm.  in  breadth;  genital 
pores  unilateral;  about  fifty  testes ;  the  uterus  consists  of  a  number  of 

'  [The  laival  stage  of  the  Davaineas  occurs  in  slugs  [Limax]  and  snails  {Helix). ~Y.  V.  T.] 


330 


THE   ANIMAL   PARASITES   OF   MAN 


loops,  which  at  each  side  are  rolled  up  into  an  almost  spherical  ball ; 
when  filled  with  eggs  the  convolutions  unwind,  permeate  the  segment 
and  then  lose  their  wall ;  the  eggs  lying  free  in  the  parenchyma 
become  finally  surrounded,  one,  or  several  together,  by  proliferating 
parenchymatous  cells ;  this  is  how  the  300  to  400  egg  masses,  taking 
up  the  entire  mature  segment,  are  formed.  The  globular  oncosphere 
(8  fi)  is  surrounded  by  two  perfectly  transparent  shells,  the  outer  of 
which  terminates  in  two  pointed  processes. 

Davainea  madagascaricnsis  has  hitherto  been  found  in  man  only 
(eight  times).  Davaine  described  this  species  from  fragments  sent 
to  him  from  Mayotta  (Comoro  Islands),  which  were  found  in  two 
Creole  children.  Chevreau  observed  four  cases  in  Port  Louis 
(Mauritius),  likewise  in  children ;  Leuckart  received  the  first  perfect 
specimen — it  was  obtained  from  a  three  year  old  boy,  the  son 
of  a  Danish  captain,  in  Bangkok  ;  Daniels,  at  the  post-mortem  of 
an  adult  native  of  George  Town,  Guiana,  found 
two  specimens  {Tcenia  dernerariensis)  ;  and  finally 
Blanchard  describes  another  perfect  specimen 
which  was  in  Davaine's  collection  of  helminthes 
in  Paris,  and  which  was  obtained  from  a  little  girl 
3  years  old,  of  Nossi-Be  (Madagascar).  The  inter- 
mediate host  is  unknown. 

Davainea  (?)  asiatica,  v.  Linst.,  1901. 

Syn.  :   Tcenia  asiatica,  v.  Linstow. 

There  exists  only  one  headless  specimen  of  this 
species,  which  is  not  quite  adult,  and  which  is  pre- 
served in  the  Zoological  Museum  of  the  Imperial 
Academy  of  Science  in  Petrograd.  It  came 
from  a  human  being  and  was  found  by  Anger  in 
Aschabad  (Asiatic  Russia,  near  the  northern  frontier 
of  Persia).  The  specimen  measures  298  mm.  in 
length.  The  breadth  anteriorly  is  only  o*i6  mm., 
the  posterior  part  measures  178  mm.  across.  The 
number  of  segments  is  about  750.  The  genita 
pores  are  unilateral ;  the  testes  are  globular  and 
lie  in  a  dorsal  and  ventral  layer  in  the  medullary  layer;  the  cirrus 
pouch  is  pyriform,  0*079  mm.  in  length  and  0*049  mm.  in  breadth  ; 
the  female  glands  lie  in  the  fore-part  of  the  segments,  the  ovary 
reaching  to  the  excretory  vessels ;  the  vitellarium  is  small  and  round. 
The  vagina  has  a  large  fusiform  receptaculum  seminis ;  the  uterus 
breaks  up  into  sixty  to  seventy  large,  irregularly  polyhedric  eggsacs. 


Fig.  234. — Scolex 
of  Davainea  niada- 
gascariensis.  The 
hooks  have  fallen 
off.  14/1.  (After 
Blanchard.) 


T^NIID^  331 

Family.     Taeniidae,  Ludwig,   1886. 
Genus.     Taenia,  L.,  1758.^ 

With  the  characters  of  the  family.  In  the  genus  Cladotaenia  recognized  by 
some  authors,  the  testes  encroach  on  the  mid  field  and  the  uterine  stem  reaches  the 
anterior  end  of  the  segment. 


Taenia  solium,  L.,  p.  p.,  1767. 

vSyn.  :  Tcenia  cucurbitina^  Pall.,  1781  ;  Tcenta  pelluctda,  Goeze,  1782;  Tcenia 
vulgaris^  Werner,  1782;  Tcenia  dentata,  Gmel.,  1790;  Haly sis  solium^  Zeder,  1800; 
Tcenia  humana  arrnata^  Brera,  1802;  Tcenia  {Cystotcenid)  solium,  Lkt.,  1862. 

The  average  lengtH  of  the  entire  tapeworm  is  about  2  to  3  m., 
but  may  be  even  more ;  the  head  is  globular,  o-6  to  o'8  to  i*o  mm. 
in  diameter.  The  rostellum  is  short  with  a  double  circlet  of  hooks, 
twenty-two  to  thirty-two  in  number,  usually  twenty-six  to  twenty-eight ; 
large  and  small  hooks  alternate  regularly;  the  length  of  the  large  hooks 
is  o'i6  to  0'i8  mm.,  of  the  small  ones  O'li  to  0*14  mm.  The  ros- 
tellum is  sometimes  pigmented.  The  suckers  are  hemispherical,  0*4  to 
o'5  mm.  in  diameter.  The  neck  is  fairly  thin  and  long  (5  to  10  mm.). 
The  proglottids,  the  number  of  which  averages  from  800  to  900,  increase 
in  size  very  gradually ;  at  about  i  m.  behind  the  head  they  are  square 
and  have  the  genitalia  fully  developed.  Segments  sufficiently  mature  for 
detachment  measure  10  to  12  mm.  in  length  by  5  to  6  mm.  in  breadth. 
The  genital  pores  alternate  fairly  evenly  at  the  lateral  margin  a  little 
behind  the  middle  of  each  segment.  The  fully  developed  uterus  con- 
sists of  a  median  trunk,  with  seven  to  ten  lateral  branches  at  either  side, 
some  of  which  are  again  ramified.  The  eggs  are  oval,  the  egg-shell 
very  thin  and  delicate  ;  the  embryonal  shell  (embryophore)  is  thick, 
with  radial  stripes ;  it  is  of  a  pale  yellowish  colour,  globular,  and 
measures  31  //,  to  36  /a  in  diameter  ;  the  oncospheres,  with  six  hooks, 
are   likewise  globular,  and  measure   20  /^  in   diameter  (fig.  238). 


'  The  Greeks  termed  the  tapeworms  eXfiivOes  irXarfTai,  more  rarely  x'/P*«  ( =  fascia) ; 
the  Romans  called  them  tienia,  tinea,  taniolay  later  lumbrici,  usually  with  the  addition  laii, 
to  distinguish  them  from  the  Luf?ibrici  teretes  (Ascarida).  The  proglottids  were  called 
Vermes  cucurbitini ;  the  cysticerci  xa^«C«'  (hailstones),  later  hydatids.  Plater  (1602)  was  the 
first  to  differentiate  Tcenia  intestinorum  {  =  Bothriocephalus  latus)  amongst  the  Lumbrici  lati 
of  man  from  Tcenia  longissima  (=  Tcenia  solium).  The  term  solitwi  was  already  used  by 
Arnoldus  Villanovanus,  who  lived, about  1300  ;  and,  according  to  him,  it  signifies  "cingulum" 
(belt,  chain),  while  N.  Andry,  in  1700,  traces  this  word  from  "solus,"  because  the  worm 
occurs  always  singly  in  man.  Leuckart  and  Krehl  derive  the  word  "  solium  "  from  the  Syrian 
"schuschl"  (the  chain),  which  in  Arabian  has  become  *' susl"  or  *' sosl,"  and  in  Latin  has 
become  "sol-ium."  What  Linnaeus  described  under  the  term  Tcenia  solium  was  really 
Tcenia  saginata  ;  the  latter  was  first  distinguished  by  Goeze,  but  was  forgotten  until  Kuchen- 
meister,  in  1852,  again  called  attention  to  the  differences. 


332 


THE   ANIMAL   PARASITES   OF    MAN 


Malformations  are  not  so  common  as  in  T.  saginaiaj  they  consist  in  two  or 
several  proglottids  being  partly  or  entirely  fused,  formation  of  single  club-shaped 
segments,  fenestration  of  long  or  short  series  of  segments  and  so-called  double  forma- 
tion, in  which  the  head  has  six  suckers  and  the  segments  exhibit  a  Y-shaped  trans- 
verse section.  The  oncospheres  occasionally  also  possess  more  than  six  booklets. 
Very  slender  specimens  have  led  to  the  description  of  a  particular  species  or  variety 
{T.  tenella). 

In  its  fully  developed  condition  T,  solium  is  found  exclusively 
in  man ;  the  head  is  usually  attached  in  the  anterior  third  of  the 
small  intestine  and  the  chain,  in  numerous  convolutions,  extends 
backwards ;  a  few  mature  detached  proglottids  usually  lie  at  the 
most  posterior  part,  and  these  are  usually  evacuated  during  defaeca- 
tion.  In  exceptional  cases  single  proglottids  or  whole  worms  may 
reach    contiguous  organs   if    abnormal    communications  with   them 


Fig.  235. — Two  fairly  mature  proglottids 
of  Tama  solium,  showing  ovaries  (one 
bi-lobed),  vitellaria,  central  uterine  stem, 
cirrus  and  vas  deferens  (above),  vagina 
(below),  testes  (scattered),  longitudinal  and 
transverse  excretory  vessels. 


Fig.  236. — Head  of  Tania  solium. 
4S/I. 


exist;  thus  they  may  reach  the  abdominal  cavity  and  the  urinary 
bladder,  or  they  may  be  found  in  a  so-called  worm  abscess  of  the 
peritoneum  ;  occasionally,  in  vomiting,  single  segments  or  several 
together  may  be  brought  up.  Exceptionally  it  induces  severe  anaemia. 
The  larval  stage  (Cysticerciis  celhilosce)  that  gives  rise  to  Tcenia 
solium  lives  normally  in  the  intramuscular  connective  tissue  and 
other  organs  of  the  domestic  pig,  but  it  is  known  to  exist  also  in 
a  few  other  mammals,  such  as  the  wild  boar,  the  sheep/  the  stag, 
dog,  cat,  brown  bear  and  monkey,  as  well  as  in  man.    The  cysticercus 

'  The  larvae  which  on  rare  occasions  are  found  in  the  muscular  system  of  sheep  are  either 
strayed  specimens  of  Cysticercus  tenuicollis^  which  normally  develop  in  organs  of  the 
abdominal  cavity,  and  appertain  to  Ti^nia  marginata  of  the  dog,  or  actually  Cysticercus 
<ellulosa.     {Cf.  Bongert,  in  Zeitschr.  f.  Fleisch-  u.  Milchhyg.,  1899,  ix,  p.  86.) 


T^NIA   SOLIUM 


333 


of  the  pig  is  an  elliptical  vesicle  with  a  longitudinal  diameter  of 
6  to  20  mm.,  and  a  transverse  diameter   of  5    to  10  mm. 

Even  with  the  naked  eye  a  white  spot  may  be  observed  in  the 
centre  of  the  long  equator,  this  being  the  invaginated  head  ;  it  is  easy 
to  make  it  project  by  pressing  on  the  vesicle  (after  tearing  off  with  the 
finger-nail  the  investing  connective  tissue),  and  on  examining  it  under 
the  microscope  one  can  convince  oneself  that  it  corresponds  with  the 
head  of  Tcenia  solium. 

Numerous  experiments  have  proved  that  the  Cysticerciis  celluloses 
of  the  pig,  if  introduced  into  the  intestine  of  man,  grows  to  a  Tcenia 
solium  (Kiichenmeister,  1855;  Humbert,  1856;  Leuckart,  1856;  Hollen- 
bach,  1859 ;  Heller,  1876) ;  the  cysticercus  has  frequently  also  been 


Fig.  237. — Large  and  small  hooks  of  Tcenia  solium.  280/] 
(After  Leuckart.) 


Fig.  238.— 7c?«m  solium.     21,  Egg  with    external   mem- 
brane ;    22,  without  (embryophore).     (After  Leuckart.) 


Fig.  239.  —  Two 
mature  proglottids 
of  Tcenia  solium  with 
fully  develope'd 
uterus.     2/1. 


cultivated  purposely  by  feeding  pigs  with  mature  proglottids  of 
T,  solium  (P.  J.  van  Beneden,  1853  ;  Haubner  and  Kuchenmeister, 
1855  ;  Leuckart,  1856  ;  Mosler,  1865  ;  Gerlach,  1870 ;  etc.),  but  success 
did  not  attend  the  efforts  to  make  Cysticercus  cellulosce  establish  them- 
selves in  the  intestines  of  pigs,  dogs,  guinea-pigs,  rabbits  and  monkeys 
{Macacus  cyuomolgus),  and  so  become  adult  Taeniae  ;  the  attempts,  also, 
to  infect  dogs  with  cysticerci  by  means  of  ova  were  likewise,  as  a 
rule,  abortive.^ 

'  According  to  Gerlach  only  young  pigs  (up  to  6  months  old)  are  capable  of  infection,  and 
perhaps  the  failure  may  have  been  due  to  the  animals  chosen  for  experiment  being  of  the 
wrong  age. 


334  THE   ANIMAL   PARASITES   OF   MAN 

The  development  of  Cysticercus  cellulosce  takes  two  and  a  half  to  three  or  four 
months  ;  it  is  not  known  how  long  the  cysticerci  remain  alive  in  animals  ;  not  uncom- 
monly they  perish  at  earlier  or  later  stages,  and  become  calcified  or  caseated. 
Extracted  cysticerci  die  in  water  at  a  temperature  of  47°  to  48''  C,  in  flesh  at  normal 
temperature  they  remain  alive  for  twenty-nine  days  or  more.  On  account  of  the 
present  rapid  means  of  pickling  and  smoking  meat,  the  cysticerci  as  a  rule  are  not 
killed,  also  the  effect  of  cold  on  them  for  some  time  in  cold  chambers  of  slaughter- 
houses is  not  lethal,  but  freezing  is  fatal  (Ostertag). 

There  is  not  the  least  doubt  that  human  beings  are  almost  exclu- 
sively infected  with  Tceiiia  solium  by  eating  pork  containing  cysticerci 
in  a  condition  that  does  not  endanger  the  life  of  the  cysticerci.  The 
infection  may  likewise  be  caused  in  man  by  eating  the  infected  meat 
of  other  animals  subject  to  this  species  of  bladder  worm,  mainly,  as  a 
matter  of  fact,  deer  and  wild  boar. 

The  frequency  of  cysticerci  in  pigs'  flesh  has  considerably  decreased  since  the 
introduction  of  meat  inspection  ;  in  the  Kingdom  of  Prussia  there  was  on  an  average 
I  infected  pig  to  every  305  slaughtered  between  1876  to  1882  ;  from  1886  to  1889,  there 
was  I  to  551  ;  from  1890  to  1892,  there  was  i  to  817  ;  in  1896,  i  to  1,470  ;  and  in  1899, 
I  to  2,T02  ;  in  the  Kingdom  of  Saxony  in  1894  there  was  i  infected  pig  to  every  636  ; 
in  1895   there  was  i   to  every  2,049,  ^^^  ^^   1896  only  i  infected  pig  was  found  of 
5,886  slaughtered.     In   South   Germany  pigs  with  cysticerci  are  very  rare,  but  are 
more  frequent  in  the  eastern  provinces  of  Prussia;   in  1892  the  number  of  infected 
pigs  compared  with  the  total  slaughtered  was  as  follows  : — 

In  the  district  of  Marienwerder     ...  ...  ...  ...         i  :       28 

„  „  Oppeln...  ...  ...  ...  ...         I  :       80 

„  „  Konigsberg  ...  ...  ...  ...         i  :     108 

„  ,,  Stralsund  and  Posen         ...  ...  ...  i  :     187 

„  „  Danzig,  Frankfort  a.  O.  and  Bromberg         ...  i  :     250 

As  compared  with  the  district  of  Arnsberg...  ...  ...         i  :     865 

„  „  „  Coblenz  ...  ...  ...         i  :     975 

„  ,,  „  Diisseldorf  ...  ...  i  :  1,070 

„  „  „  Miinster  and  Wiesbaden     ...         i  :  1,900 

The  average  for  the  whole  of  Prussia  in  the  same  year  was  i  :  1,290;  for  the 
eastern  provmces,  on  the  other  hand,  i  :  604.  Even  more  unfavourable  are  the  pro- 
portions in  Russian  Poland  (over  i  per  cent,  of  pigs  measly),  in  Prague  (over  3  per 
cent.),  in  Bosnia  and  Herzegovina  (6  to  7  per  cent.).  The  cause  for  this  is  most 
likely  attributable  to  the  manner  in  which  the  pigs  are  kept.  When  allowed  to  be 
in  the  farmyards  of  the  small  farmers  for  the  whole  day,  or  allowed  to  wander  in  the 
village  streets  and  pasture  lands,  they  are  more  liable  to  take  up  the  oncospheres  of 
the  T.  solium  than  when  shut  up  in  good  pig-styes. 

The  geographical  distribution  of  T.  solium  generally  corresponds 
with  that  of  the  domestic  pig  and  the  custom  of  eating  pork  in  any 
form  insufficiently  cooked  or  raw.  There  are,  or  were,  some  isolated 
districts  in  Germany,  France,  Italy  and  England  where  the  "armed 
tapeworm"  was  frequent  (for  instance,  Thuringia,  Brunswick,  Saxony, 
Hesse,  Westphalia,  whereas  it  is  and  was  very  scarce  in  South  Ger- 
many) ;  it  is  thus  easily  understood  why  it  occurs  very  rarely  in  the 
East,  in  Asia  and  in  Africa,  in   consequence  of  the  Mahommedans, 


T^NIA   SOLIUM  335 

Jews,  etc.,  not  eating  pork.  In  North  America,  also,  T.  solium  is  very 
rare  ;  the  tapeworm  which  is  known  there  by  this  name  is  generally 
T.  sagiiiata,  Stiles.  During  the  last  decade  T.  solium  infection  has, 
however,  very  markedly  decreased  in  North  and  East  Germany  in 
consequence  of  the  precautions  exercised  by  the  public  in  the  choice 
of  pork  to  avoid  trichinosis,  especially,  however,  because  measly  meat 
must  be  sold  as  such  and  must  be  thoroughly  cooked  before  being 
placed  on  the  market ;  indeed,  if  badly  infected  it  may  not  be  sold 
for  food,  but  can  be  turned  to  account  for  industrial  purposes. 

The  occurrence  of  Cysticercus  cellulosce  in  man  has  been  known 
since  1558  (Rumler,  Obs.  med.,  liii,  p.  32);  there  is  hardly  an  organ 
in  man  in  which  cystercerci  have  not  been  observed  at  some  time  ; 
they  are  most  frequently  found  in  the  brain,^  where  they  grow  to  a 
variety  known  as  Cysticerciis  racemosus ;  next  in  frequency  they  are 
found  in  the  eye,  in  the  muscular  system,  in  the  heart,  in  the  sub- 
cutaneous connective  tissue,  the  liver,  lungs,  abdominal  cavity,  etc. 
The  number  of  cysticerci  observed  in  one  man  varies  between  a  few 
and  several  thousands.  Of  the  sexes,  men  are  most  subject  (60  to 
66  per  cent,  of  the  number  attacked).  The  disturbances  caused  in 
man  by  cysticerci  vary  according  to  the  nature  or  position  of  the 
organs  attacked  ;  when  situated  in  the  cerebral  meninges  they  have 
the  same  effect  as  tumours. 

During  the  last  decades,  however,  these  cases  have  also  become 
less  common.  In  Rudolphi's  time  2  per  cent,  of  post-mortems  in 
Berlin  showed  cysticerci  ;  in  the  'sixties,  according  to  Virchow,  about 
the  same  ;  in  1875  the  number  fell  to  1*6  per  cent ;  in  1881  to  0*5  per 
cent.  ;  in  1882  to  0*2  per  cent. ;  in  1900  to  0*15  per  cent.,  and  in  1903 
to  o'i6  per  cent.  Hirschberg  between  1869  and  1885  discovered 
cysticerci  in  the  eye  seventy  times  in  60,000  ophthalmic  cases,  but 
during  the  following  six  years  the  parasite  was  only  present  twice 
amongst  a  total  of  46,000  cases  of  ophthalmic  diseases,  and  since 
1895  no  ophthalmic  case  has  been  met  with. 

The  infection  of  human  beings  with  the  cysticerci  can  only  take 
place  by  the  introduction  of  the  oncospheres  of  Tcenia  solium  into 
the  stomach  with  vegetable  foods,  salads  that  have  been  washed 
in  impure  water  containing  oncospheres,  also  by  drinking  con- 
taminated water  ;  the  carriers  of  T.  solium,  moreover,  infect  themselves 
still  more  frequently  through  uncleanliness  in  defaecation,  the  privies 

'  Dressel,  for  instance,  examined  eighty  seven  persons  suffering  from  cysticercus,  and 
found  it  seventy-two  times  in  the  brain,  thirteen  times  in  the  muscles  ;  K.  Muller,  in  thirty- 
six  cases,  found  it  twenty-one  times  in  the  brain,  twelve  times  in  the  muscles,  three  times  in 
the  heart ;  Haugg,  in  twenty-five  cases,  found  it  thirteen  times  in  the  brain,  six  times  in  the 
muscles,  twice  in  the  skin,  etc.  According  to  Graefe,  amongst  i,ooo  ophthalmic  cases  in 
Halle  and  Berlin,  there  was  one  with  cysticercus  in  the  eye ;  in  Stuttgart  there  was  only  one 
in  4,000,  in  Paris  one  in  6,000,  and  in  Copenhagen  one  in  8,000. 


336  THE  ANIMAL   PARASITES   OF   MAN 

in  public  localities  and  many  private  houses  affording  striking  testi- 
mony of  this.  The  minute  oncospheres  can  thus  easily  reach  the 
fingers  and  thence  the  mouth  (as  in  twirling  the  moustache,  biting  the 
nail).  More  rarely  a  third  manner  of  transmission  or  internal  auto- 
infection  may  possibly  take  place,  as  when,  in  the  act  of  vomiting, 
mature  proglottids  near  the  stomach  are  drawn  into  it  ;  the  onco- 
spheres or  segments  there  retained  are  then  in  the  same  position  as 
if  they  had  been  introduced  through  the  mouth. 

On  account  of  these  dangers  of  internal  or  external  auto-infection,  it  is  therefore 
the  duty  of  the  medical  attendant,  after  recognizing  the  presence  of  tapeworms,  to 
expel  them,^  and  in  doing  so  to  employ  every  possible  means  to  prevent  vomiting 
setting  in  ;  it  is,  however,  equally  important  to  take  the  necessary  steps  to  destroy 
the  parasites  evacuated.  It  may  be  incidentally  mentioned  that  in  using  certain 
remedies  the  scolex  not  rarely  remains  in  the  intestine  ;  the  cure  in  such  cases  has 
not  been  accomplished,  as  the  scolex  again  produces  new  proglottids,  and  after 
about  eleven  weeks  the  first  formed  ones  are  again  mature  and  appear  in  the  fasces. 

Amongst  the  cysticerci  also  many  malformations  appear ;  thus  absence  of  the 
rostellum  and  the  hooks,  or  double  formation  with  six  suckers,  or  abnormalities  of 
growth  on  account  of  the  surroundings,  which  have  had  a  special  name  given  to 
them,  viz.,  Cysticercus  racernosus^  Zenk.  {=  C.  botryoides,  Hell.  ;  C.  multtlocularisj 
Kchnmstr.) ;  these  forms  are  more  especially  found  at  the  base  of  the  brain,  are 
irregularly  ramified  and  often  without  the  head. 

A  certain  interest  is  attached  to  those  forms  that  have  led  to  the 
establishment  of  a  distinct  species  : — 


Cysticercus  acanthotrias,  Weinld.,  1858. 

In  making  the  autopsy  of  a  white  Virginian  woman  who  had  died  of  phthisis,  a 
cysticercus  was  found  in  the  dura  mater,  and  eleven  or  twelve  specimens  in  the 
muscles  and  subcutaneous  tissue.  Weinland  and  Leuckart,  who  examined  the 
specimens,  found  that  they  resembled  Cysticercus  celluloscB  in  form  and  size,  but  that 
they  carried  on  the  rostellum  a  triple  crown,  each  consisting  of  fourteen  to  sixteen 
hooks,  which  differed  from  the  hooks  of  C.  celluloses  or  of  Tcenia  solium  by  the 
greater  length  of  the  posterior  root  process  and  the  more  slender  form  of  the 
hooks;  the  large  hooks  measured  0-153  to  0-196  mm.,  the  medium-sized  hooks, 
o'l  14  to  0-14  mm.,  and  the  small  ones  0-063  to  0-07  mm. 

On  account  of  these  differences  a  distinct  species  of  cysticercus 
was  established,  and  this  naturally  presupposed  a  corresponding 
species  of   Taenia  (7.  acanthotrias,   Lkt.)  ;    this  could  be   done  with 

'  The  diagnoiss  as  a  rule  is  not  difficult ;  the  patients  themselves  frequently  observe  the 
pumpkin  seed-like  segments  in  the  faeces.  But  in  such  cases  the  diagnosis  must  still  be 
confirmed.  In  other  cases  the  discovery  of  the  oncospheres  in  their  embryonal  shells 
(embryophores),  which  cannot  be  confounded  with  the  other  constituents  of  the  faeces,  gives 
complete  certainty,  although  the  dififerential  diagnosis  between  T.  solium  and  T.  raginata 
is  hardly  possible  from  the  embryophoies  ;  but,  if  evacuated  segments  are  placed  between 
two  slides  and  lightly  pressed,  the  species  is  easily  recognizable  by  the  shape  of  the  uterus 
{cf.  figs.  239  and  241). 


T^NIA   BREMNERI  337 

justice  so  long  as  the  case  remained  isolated,  Le.j  in  America,  as 
there  was  the  possibihty  of  the  corresponding  Taenia  being  found.  In 
this  respect,  however,  the  position  has  changed  ;  Delore  first  described 
a  cysticercus  the  size  of  a  nut  from  the  biceps  muscle  of  the  arm  of 
a  silk-worker  in  Lyons  ;  according  to  Bertolis  this  specimen  possessed 
hooks  of  three  different  sizes,  the  dimensions  of  which  corresponded 
with  the  figures  given  by  Weinland  and  Leuckart ;  the  correctness  of 
the  diagnosis  could  hardly  be  doubted,  as  Bertolis  was  known  to  be 
a  very  exact  observer.  A  second  case  has  become  known  through 
Cobbold,  who  regards  a  specimen  of  a  cysticercus  in  Dallinger's 
collection  as  likewise  belonging  to  Cysticercus  acanthotrias ;  this 
specimen  also  came  from  a  man's  brain ;  finally  a  third  case,  also 
from  France,  has  been  published  by  Redon.  This  author,  amongst 
numerous  C.  celluloscv  of  a  man,  found  one  that  had  forty-one  hooks  in 
three  rows,  and  he  was  the  first  to  express  the  opinion  that  C.  acantho- 
trias  does  not  represent  a  distinct  species,  but  is  only  an  abnormality 
of  C.  cellulosce.  This  view  was  also  taken  by  Blanchard  and  Railliet,  and 
is  probably  correct,  as  the  discovery  of  the  large  corresponding  Taenia 
furnished  with  three  rows  of  hooks  is  not  to  be  expected  in  European 
beasts  of  prey,  and  in  Redon's  case  C.  acanthotrias  as  well  as  C.  cellulosce 
occurred  simultaneously. 

The  duration  of  life  of  C.  cellulosce  in  man  is  very  long ;  cysticerci 
of  the  eye  have  been  known  to  persist  for  twenty  years,  and  in  cysti- 
cercus of  the  brain  ten  to  nineteen  years  may  elapse  from  the  first 
appearance  of  cerebral  symptoms  until  death.  Dead  cysticerci  may 
shrivel  up  or  become  calcified,  perhaps  also  undergo  fatty  degeneration 
and  then  absorption.  Finally,  it  may  be  mentioned  that  if  particular 
proof  is  required  that  C.  cellulosce  of  man  belongs  to  the  cycle  of 
development  of  the  Tcenia  solium,  such  proof  has  been  furnished  by 
Redon. 

Note. —  Tcenia  tenella,  mentioned  on  p.  332,  was  ascribed  by  Cobbold  to 
cysticerci  of  the  muscular  system  of  sheep.  It  has,  however,  been  demonstrated 
that  these  cysticerci  belong  to  the  cycle  of  development  of  Tcenia  marginata 
(dog)  {Cysticercus  tenuicollis^  from  the  omentum  of  sheep)  ;  but  as  already  stated  ^ 
C.  cellulosce  also  occurs  in  sheep.  Chatin  himself  swallowed  the  cysticercus,  which 
Cobbold  termed  C.  ovis,  without  causing  a  Taenia  to  develop  in  his  intestine.  Miiller 
also  vainly  sought  to  induce  infection  with  C.  tenuicollis  in  his  own  person.  On  the 
other  hand,  the  feeding  of  a  dog  with  Cysticercus  ovis  resulted  in  the  production  of 
Tcenia  marginata. 

Taenia  bremneri,  Stephens,  1908. 

Characterized    by  the    large   size    of   the    gravid    segments.      The 
largest  was  32  by  9  mm.     Smallest  21   by  6  mm.     Average  28*6  by 
8*5    mm.      Mode    21    by   6  mm.      Uterine   branches  twenty-two  to 
22 


338 


THE    ANIMAL   PARASITES   OF   MAN 


twenty- four  in  number.  Calcareous  bodies  numerous,  15-2  /x  in 
diameter.  Eggs  maximum  45*6 /t  by  41-8 /-t.  Smallest  34*2  ^la  by 
30-4  yLt.     Mode  38 /x  by  30*4 /a. 

Taenia  marginata,  Batsch,   1786. 
Syn. :  T.  e.  Cysticerco  ienuicolH,  Kiichenmeister,  1853. 
This  species,  which  in  structure  resembles  Tcenia  solium^  lives  in  the  intestine 
of  the  dog  and  the  wolf.      It  attains  1*5  to  4  m.  in  length,  possesses  a  double  crown 

of  thirty  to  forty  hooks,  on 
an  average  thirty-six  to  thirty- 
eight  hooks,  and  in  its  larval 
stage  {Cysticercus  tenuicollis) 
lives  in  the  peritoneal  cavity  of 
ruminants  and  the  pig,  occasion- 
ally in  the  monkey  and  squirrel. 
It  is  included  in  this  work 
because,  according  to  one  state- 
ment, C.  tenuicollis  is  supposed 
to  have  been  observed  in  man 
in  North  America  ;  but  the  case 
is  not  quite  certain,  as  the  number  of  hooks  was  less  than  in  C.  tefiuicollis  and 
coincided  with  C.  cellulosce^  although  the  size  of  the  cysticercus  appeared  to  point  to 
C.  ienuicolHs.  A  yet  earlier  statement  of  Eschricht,  that  Cysticercus  tenuicollis  had 
been  observed  in  Iceland  in  the  liver  of  a  man,  is  undoubtedly  due  to  an  error. 


Fig.  240. — Large  and  small  booklets  of  Tania 
marginata.     280/1.     (After  Leuckart.) 


Taenia  serrata,  Goeze,  1782. 

This  parasite  attains  a  length  of  from  0*5  to  2  m.,  possesses  a  double  crown  of 
thiity-four  to  forty-eight  (mostly  forty)  hooks.  It  lives  exclusively  in  the  intestine 
of  the  dog,  the  corresponding  cysticercus  {Cysticercus  pisiforinis)  living  in  the 
mesentery  of  the  hare  and  rabbit.  We  mention  this  species  with  all  reserve 
amongst  the  parasites  of  man,  because  Vital  states  that  he  has  observed  it  twice  in 
Constantine  (Algeria)  in  human  beings.  The  data,  however,  are  not  sufficient  to 
characterize  the  species.  It  is  highly  probable  that  they  relate  to  Tcenia  solium. 
Galli-Valerio  even  swallowed  five  specimens  of  Cysticercus  pisiformis^  but  without 
result. 

Taenia  crassicollis,  Rud.,   1810. 

I  only  mention  this  species  from  the  intestine  of  the  domestic  cat  because 
Krabbe  regards  its  occurrence  in  man  as  possible.  It  attains  a  length  of  60  cm. 
and  is  armed  ;  its  cysticercus  {Cysticercus  fasciolaris)  lives  in  the  liver  of  mice  and 
rats.  In  Jutland,  according  to  Krabbe,  chopped-up  mice  (spread  on  bread)  are 
eaten  raw,  being  a  national  remedy  for  retention  of  urine,  and  this  custom  affords 
the  possibility  of  the  introduction  of  C.  fasciolaris  into  the  intestine  of  man  {Nord. 
nied.  Arkiv,  1880,  xii). 

Taenia  saginata,  Goeze,  1782. 

Syn.  :  Tcenia  solium^  L.,  1767  {pro parte)  ;  Tcenia  cucurbitina,  Pallas,  1781  [p.p.)  ; 
Tcsnia  inertnis^  Brera,  1802.  Moquin-Tandon,  i860;  Tcenia  dentata,  Nicolai,  1830; 
Tcenia  lata.,  Pruner,  1847;  Bothriocephalis  tropicus^  Schmidtmuller,  1847;  Tcenia 
mediocanellata,  Kiichenmeister,  1855  ;  Tcenia  zittavensis^  Kiichenmeister,  1855  ;  Tcenia 
tropica^  Moquin-Tandon,  i860;   Tcenia  {Cystotcenia)  mediocanellata.,  Leuckart,  1863. 


T^NIA    SAGINATA 


339 


The  length  of  the  entire  tapeworm  averages  4  to  8  to  10  m.  and 
more,  even  up  to  36  m.  According  to  Berenger-Feraud  it  attains 
a  length  of  74  m.  (?)  The  head  is  cubical  in  shape,  1*5  to  2  mm. 
in  diameter  ;  the  suckers  are  hemispherical  (o*8  mm.)  and  are 
frequently  pigmented  ;  there  is  a  sucker-like  organ  in  place  of  the 
rostellum,  and  this  also  is  frequently  pigmented.  The  neck  is 
moderately  long  and  about  half  the  breadth  of  the  head  ;  the 
proglottids,  the  number  of  which  averages  more  than  1,000, 
gradually  increase  in  size  ;  the  mature  detached  segments  are  shaped 
exactly  like  pumpkin-seeds,  and  are  about  16  to  20  mm.  in  length 
and  4  to  7  mm.  in  breadth.  The  genital  pores  alternate  irregularly 
and  are  situated  somewhat  behind  the  middle  of  the  lateral  margin. 

There  are  twenty  to  thirty-five 
lateral  branches  at  each  side  of  the 
median  trunk  of  the  uterus,  and 
these  again  ramify.  The  eggs  are 
more  or  less  globular,  the  egg- 
shell frequently  remains  intact  and 
carries  one  or  two  filaments  ;    the 


Fig.  241. — Mature 
segment  of  Tcenia 
saginata,  G.,  with 
distended     uterus. 

2/1. 


Fig.  242.  —  Ce- 
phalic end  of  Tania 
saginata  in  the  con- 
tracted condition. 
8/1. 


Fig.  243. — Tania  saginata.  19,  egg  with 
external  shell.  20,  without  (embryophore), 
(After  Leuckart.) 


embryonal  shell  (embryophore)  is  thick,  radially  striated,  is  transparent 
and  oval  ;  it  is  30  jx  io  40  /a  in  length,  and  20  /^  to  30  /^  in  breadth. 
Several  segments  simultaneously  are  usually  passed  spontaneously 
with  defaecation. 


Malformations  are  not  uncommon,  and  resemble  those  of  Tcenia  solium; 
a  triangular  form  has  been  termed  T.  capensis  by  Kiichenmeister,  and  T.  lopho- 
soma  by  Cobbold,  names  that  naturally  possess  as  little  value  as  does  the  term 
T.  fenestrata  for  fenestrated  specimens.  Moreover,  T.  solium,  var.  adietina,  Weinld., 
1858,  which  was  evacuated  by  an  Indian,  was  probably  a  T.  saginata  with  somewhat 
close  uterine  branches.  It  is  regarded  by  Stiles  and  Goldberger  as  a  doubtful  sub- 
species. 

r.  saginata  in  its  adult  condition  lives  exclusively  m  the  intestinal 


340 


THE   ANIMAL   PARASITES   OF   MAN 


canal  of  man.^  The  corresponding  cysticerus  is  Cysticercus  hovis,  and 
is  found  almost  exclusively  in  the  ox  ;  it  is  small,  7*5  ^o  9  '^^^ 
in  length  and  5*5  mm.  in  breadth,  may  easily  escape  notice,  and 
requires  from  three  to  six  months  for  its  development.  Numerous 
experiments  have  confirmed  the  connection  of  Cysticercus  bovis  with 
Tcenia  saginata  ;  indeed,  the  cysticercus  was  only  discovered  by  feeding 
experiments  after  attention  had  been  called  to  the  ox  as  the  probable 
intermediary  host  of  this  Taenia. 

Medical  men  observed  that  weakly  children  who  were  ordered  to  eat  raw  scraped 
beef  to  strengthen  them  contracted  T.  saginata.  It  was  found,  moreover,  thai 
Jews,  who  are  prohibited  from  eating  pork  from  religious  motives,  suffered  especially 
from  T.  saginata;  when  T.  solium  was  found  to  occur  in  a  Jew  he  often  con- 
fessed to  having  eaten  pork  ;  and  finally  it  was  found  that  certain  nations — for 
instance,  the  Abyssinians— frequently  harbour  T.  saginata,  and  only  eat  beef— raw 
by  preference. 

These  observations  led  Leuckart,  in  1861,  to  feed 
young  calves  with  the  proglottids  of  T.  saginata  in 
order  to  discover  the  corresponding  cysticercus,  which 
was  then  not  known.  This  experiment  was  successful. 
Similar  experiments,  with  similar  results,  were  then  con- 
ducted by  Mosler  (1863),  Cobbold  and  Simonds  (1864 
and  1872),  Roll  (1865),  Gerlach  (1870),  Zurn  (1872), 
Saint  Cyr,  Jolicceur  (1873),  Masse  and  Pourquier 
(1876),  and  Perroncito,  in  1876.  The  attempts  to 
infect  goats,  sheep,  dogs,  pigs,  rabbits  and  monkeys 
were  unsuccessful.  Only  Zenker  and  Heller  were  able 
to  infect  kids,  and  Heller  infected  one  sheep,  but  these 
are  exceptions. 

Artificial  infections  of  human  beings  with 
Cysticercus  bovis  to  obtain  the  tapeworm  were 
less  numerous,  and  indeed  quite  superfluous, 
yet  this  was  also  done  by  Oliver  (1869)  in  India, 
and  Perroncito  (1877)  in  Italy.  The  experi- 
ments of  the  latter  prove  that  the  extracted 
cysticerci  of  the  ox  certainly  perish  in  water  at 
47°  to  48°  C. 


Fig.  244. — A  piece  of 
the  muscle  of  the  ox,  with 
three  specimens  of  Cysticer- 
cus bovis.  Natural  size. 
(After  Ostertag.) 


It  is  a  remarkable  circumstance  that,  at  least  as 
regards  Central  Europe,  C.  bovis  in  the  ox,  after  natural 
infection,  was  so  seldom  found  that  almost  every  case 
was  published  as  a  rarity ;  whereas  the  Taenia  is  very  frequent  in  man.  The 
reason  for  this  is  that  in  Germany  cattle  are  not  severely  infected,  and  that  the 
small,  easily  dried-up  cysticerci  easily  escape  notice  in  the  large  body  of  the  host. 
Hertwig,  the  late  director  of  the  town  cattle  market  in  Berlin,  in   1888,  pointed   out 


•  Abnormal  migrations  of  this  species  have  also  been  known.  Compare,  amongst 
others,  Stieda,  A.,  '*  Durchbohr.  d.  Duod.  u.  d.  Pancreas  durch  eine  Taenia,"  Centralbl. 
f.  Bakt.,  Path,  und  Infektionsk.,  1900,  xxviii  (i),  p.  430. 


T^NIA   SAGINATA 


341 


that  the  cysticercus  of  the  ox  is  found  chiefly  in  the  musculi  pterygo.de  externi 
and  interni,  and  since  that  time  a  far  greater  number  of  infected  oxen  have  been 
found  in  Berlin. 


Year 

Number  of  oxen 
slaughtered 

Infected 

Proportion 

1888—89 
1889—90 
1890—91 
1891—92 
1892—93 

141,814 
154,218 

124,593 
136,368 
142,874 

113 
390 
263 
252 
214 

I  :  1,255 
I  :      395 
I  :     474 
I  :      541 
I  :      672 

Since  1892  an  increase  has  taken  place  in  the  number  of  oxen  infected  with 
cysticercus,  but  this  is  probably  attributable  to  the  more  general  and  searching 
examinations.  In  the  slaughter-houses  of  Prussia  the  number  of  infected  beasts  was 
as  follows  : — 


1892 

567 

1893 

686 

1894 

748 

1895 

1,143 

1896 

...         1,981 

1897 

2,629 

The  condition  was  most  frequent  in  Neisse  (3*2  to  4  per  cent.),  Eisenach 
(I'Qi  per  cent.),  Ohlau  (1*57  per  cent.),  Oels  i.  Schles.  (i  per  cent.),  Marienwerder 
(o'34  to  I '2  per  cent.).  The  flesh  of  oxen  only  slightly  infected  (containing  not  more 
than  ten  living  cysticerci)  is  sold  in  pieces  of  not  more  than  5  lb.  to  consumers 
after  having  been  rendered  innocuous  by  cooking,  or  by  pickling  for  twenty-one  days 
in  25  percent,  salt  brine,  or  hanging  for  twenty-one  days  in  suitable  refrigerators; 
oxen  in  which  only  one  cysticercus  is  found  are  allowed  free  commerce,  and  those 
strongly  infected  {i.e.^  containing  more  than  ten  living  cysticerci)  may  only  be  used 
for  industrial  purposes. 

It  is  a  striking  fact  that  more  bulls  than  cows  are  infected  (according  to  Reiss- 
mann,  in  Berlin,  from  1895  to  1902,  o"446  per  cent,  bulls,  0*439  pei*  cent,  oxen,  and 
0*262  per  cent,  cows),  the  explanation  of  which,  according  to  Ostertag,  is  that  most 
oxen  are  killed  when  young,  when  also  infection  most  readily  takes  place,  and, 
further,  that  the  larva  later  on  in  life  can  be  completely  atrophied. 

The  cysticercus  of  the  ox  has  hitherto  been  found  in  man  on  very 
rare  occasions.  Arndt  {Zeitschr.  f.  Psychiat.,  xxiv)  mentions  a  case  in 
the  brain,  Heller  in  the  eye,  and  Nabiers  and  Dubreith  also  in  the 
brain  {Journ.  med.  Bordeaux,  1889 — 1890,  p.  209)  ;  but  the  diagnoses 
are  not  quite  certain,  as  absence  of  hooks  occasionally  occurs  in 
Cysticercus  cellulosce. 

Tcenia  saginata  is  the  most  frequent  tapeworm  of  man  (with  the 
exception  of  Dibothriocephalus  latus  in  a  few  districts),  and  the  parasite 
is  widely  distributed  over  the  surface  of  the  globe ;  it  has  been  known 
in  the  East  for  ages,  so  far  as  data  are  available ;  it  is  frequent  in 
Africa,  America,  and  Europe.  Its  frequency  has  perceptibly  increased 
during  the  last  few  years,  but  a  decrease  should  soon   take  place  in 


342 


THE   ANIMAL   PARASITES   OF   MAN 


consequence  of  the  extent  and  improvement  of  the  official  inspection 
of  meat. 

The  following  table  shows  the  relative  frequency  of  the  Cestodes 
of  man  : — 


Author 

Year 

Number  of 
cases 

T.  saginata 

T. 
solium 

Dibr. 
latus 

Di/>yl. 
canin. 

Unde- 
termined 

Parona  (Milan)     

1899 

150 

121 

II 

4 

— 

14 

Parona  (Italy)       

1868-99 

513 

397 

71 

26 

— 

19 

Krabbe  (Denmark) 

1869 

100 

37 

53 

9 

I 

»»               »» 

1869—86 

200 

'53 

24 

16 

8 

— 

>»               »> 

1887-95 

100 

89 

— 

5 

6 

— 

j>               »» 

1896— 1904 

50 

41 

I 

S 

3 

— 

Blanchard  (Paris) 

1895 

? 

1,000 

21 

— 

Stiles  (United  Slates)      ... 

1895 

( more  than 
1      300 
19 

more  than) 
30c    1 
16 

— 

3 

— 

— 

Schoch  (Switzerland) 

1869 

I 

2 

— 

— 

Zaeslein  (Switzerland)     ... 

1881 

? 

180 

19 

? 

— 

— 

Kessler  (Petrograd) 

1888 

? 

22 

16 

47 





Mosler  (Greifswaid) 

1894 

181 

112 

64 

5 

— 

— 

Bollinger  (Munich) 

1885 

25 

16 

8 





Vieroidt  (Tubingen) 

1885 

121 

"3 

8 

— 

— 

— 

Mangold  (Tubingen) 

1885-94 

128 

120 

6 

8 

— 

— 

Taenia  africana,  v.  Linst.,  1900. 

This  worm  measures  over  i'3  m.  in  length.  The  segments  are 
all  broader  than  they  are  long.  The  scolex  is  unarmed  and  is 
provided  with  an  apical  sucker  (o*i6  mm.).  The  parasite  measures 
1-38  mm.  in  breadth,  i'03  mm.  in  width ;  the  suckers  measure 
0*63  mm.  in  diameter.     The  neck  is  very  short  and  somewhat  broader 


Fig.  245.— Mature  segment  of  Tcenia  africana.  The  ovary  is  in  the 
middle,  and  behind  it  are  the  shell  gland  and  vitellarium  ;  at  the  sides  are 
the  testicles,  and  externally  the  excretory  canals;  the  cirrus  pouch,  the  vas 
deferens  and  the  vagina  are  on  the  left.     Magnified.      (After  v.  Linstow.) 

than  the  scolex;  number  of  segments  about  600;  the  hindmost 
segments  measure  7  mm.  in  length  and  12  to  15  mm.  in  breadth. 
The  genital  pores  alternate  irregularly  in  the  middle  of  the  lateral 
margin  ;  the  testes  are  very  numerous  and  occupy  the  entire  medul- 
lary layer;  the  vas  deferens  is  much  convoluted;   the  cirrus   pouch 


T^NIA   CONFUSA 


343 


is  pyriform  and  thick  walled  ;  the  cirrus  and  vagina  are  beset  with 
bristles  directed  outwards  ;  the  receptaculum  seminis  is  fusiform  ;  the 
ovary  is  large  and  double,  and  consists  of  radially  placed  club-shaped 
tubes  that  do  not  anastomose  and  do  not  branch :  the  vitellarium  is 


Fig.  246. — Proglottis  of  Tcenia  africana,  wiih  uterus.     Magnified. 
(After  V.  Linstow.) 

at  the  posterior  border  of  the  proglottids,  the  round  shell  gland  in 
front  of    it ;    the  uterus  consists  of   a  median  trunk  and  fifteen  to 

twenty-four  non-ramified  lateral  branches 
on  each  side  ;  the  embryonal  shell  is  thick 
and  has  radial  stripes — it  may  be  round 
(31-2//,  to  SS'B//-)  or  oval  (39/x  by  33*8/^); 
the  spines  of  the  oncospheres  measure 
7  yu,  to  8  yn  in  length  (fig.  197). 

At   present  only  two  specimens  are 
known  ;  they  came  from  a  black  soldier 
from  the  vicinity  of   Lake  Nyasa.     The 
cysticercus  is  unknown ;  perhaps  it  lives 
rnlf:  ?n5;:;^.nl5  ^^!\t£l'     i"  thc  zcbu,  thc  fiesh  of  which  thc  Askaris 

<ana ;     apical    surrace.     Magnified.  ' 

<After  V.  Linstow.)  are  in  the  habit  of  devouring  uncooked. 


Taenia  confusa,  Ward,  1896. 

Length  8*5  m.,  breadth  about  5  mm.  The  scolex  is  unknown  ; 
there  is  no  neck;  number  of  proglottids  700  to  800,  always  longer 
than  they  are  broad ;  the  hindmost  measure  35  mm.  in  length,  4  to 
5  mm.  in  breadth  ;  the  genital  pores  alternate  irregularly  behind  the 
middle  of  the  lateral  margin  ;  testicles  numerous  ;  vas  deferens  not 
much  coiled  ;  the  cirrus  pouch  thick  walled,  elongated  and  club- 
shaped,  with  globular  vesicula  seminalis;  the  cirrus  is  beset  with  little 
hairs  ;  the  receptaculum  seminis  is  globular ;  ovary  small,  double  ; 
each  half  is  bean-shaped  ;  vitellarium  narrow,  triangular  ;  shell 
gland  globular;  uterus  with  median  trunk  and  fourteen  to  eighteen 
short  ramified  lateral  branches  on  either  side.  The  embryophores  are 
oval  (39  yit  by  30  /a),  thick  and  radially  striated. 


344 


THE    ANIMAL   PARASITES    OF   MAN 


Of   this  species  only  two  specimens  have    been   recorded ;    they 
occurred  in  human   beings  and  were  sent  at  different  times  to  the 


Fig.  248.  —  Tcenia  confusa  :  mature 
segment  showing  central  uterine  stem, 
bilobed  ovary,  globular  shell  gland,  trian- 
gular vitellarium,  scattered  testes,  cirrus, 
vas  deferens,  and  vagina.  15/1.  (After 
Guyer.) 


Fig.  249.  —  TcBnia  con- 
fusa :  gravid  segment. 
25/1.     (After  Ward.) 


first  describer  of  them  by  a  doctor  in  Lincohi  (Nebr.).  Perhaps 
Tcenia  solium,  var.  abietina,  Weinld.,  which  was  found  in  a  Chipe- 
way  Indian,  is  of  the  same  species  in  spite  of  the  shorter  segments. 


Taenia  echlnococcus,  v.  Sieb.,  1853. 

Syn.  :    Tcenia  nana^  v.  Ben.,  1861  {nec\.  Sieb.,  1853)  ;    Echinococcifer 
echlnococcus,  Weinld.,  1861. 

Tcenia  echinococcus  measures  2*5  to  5  or  6  mm.  in  length  ;  the  head 
is  0'3  mm.  in  breadth,  and  has  a  double  row  of  twenty-eight  to  fifty 
booklets  (on   an  average  thirty-six  to  thirty-eight)  on  the  rostellum. 

The  size  and  form  of  these  booklets  vary  (the  larger  ones  are 
0*040  to  0*045  ^^"^-  i"  length,  the  smaller  ones  are  0*030  to 
0*038  mm.  in  length).  The  suckers  measure  0*13  mm.  in  diameter; 
the  neck  is  short ;  there  are  only  three  or  four  segments,  the  posterior 
segment  being  about  2  mm.  in  length  and  0*5  mm.  in  breadth.  The 
genital  pores  alternate  ;  there  are  forty  to  fifty  testicles  ;  the  vas  deferens 


T^NIA   ECHINOCOCCUS 


345 


is  spirally  coiled;  the  cirrus  pouch  is  pyriform.  The  ovary  is  horse- 
shoe-shaped with  the  concavity  directed  backwards;  the  vitellarium 
double,  each  half  almost  bean-shaped,  at 
right  angles  to  the  plane  of  the  segment ; 
the  shell  gland  is  round.  The  median  trunk 
of  the  uterus  is  dilated  when  filled  with  eggs 
and  (instead  of  lateral  branches)  has  lateral 
diverticula.  It  is  not  unusual  for  the  eggs 
to  form  local  heaps.  The  embryonal  shell 
(embryophore)  is  moderately  thin,  with  radial 
striae,  almost  globular,  30  />t  to  36  /x  in 
diameter. 

When  mature  this  parasite  lives  in  the 
small  intestine  of  the  domestic  dog,  the 
jackal,  and  the  wolf,  and  apparently  also  in 
Felis  concolor,  and  is  usually  present  in  great 
numbers  ;  it  can  also  be  transmitted  ex- 
perimentally to  the  domestic  cat,  one  suc- 
cessful result  out  of  seven  (Deve).^  The 
larval  stage  (Echinococcus  polymorphus)  lives 
in  various  organs — chiefly  in  the  liver  and 
lungs — of  numerous  species  of  mammals 
(twenty-seven),  especially  in  sheep,  ox  and 
pig,  and  it  is  even  not  uncommon  in  man, 
though  the  Taenia  itself  has  never  been  found 
in  a  human  being;  accordinglyman  can  only 
acquire  the  echinococcus  by  ingesting  the 
eggs  of  the  "  dog  worm."  The  dogs  dis- 
seminate the  eggs  of  Tcenia  echinococcus 
wherever  they  go,  or  carry  them  to  their 
mouths  and  coats  by  biting  up  the 
evacuated  segments,  and  are  thus  able  to 
transmit  them  directly  to  human  beings  (by 
licking  them  or  making  use  of  the  same 
crockery,  etc.).  In  other  cases  the  onco- 
spheres, enclosed  in  the  embryophores,  must 
withstand  desiccation  for  a  time  and  then 
(as  when  the  dogs  are  ''  kissed  "  or  other- 
wise caressed)  are  transmitted  into  or  on  to    50/1. 


Fig.  250.  —  Tcenia  echino- 
coccus :  the  cirrus  sac,  the 
vagina,  uterus,  ovary,  shell  gland 
and  vitellarium,  and  the  testicles 
at  the  sides  are  recognizable  in 
the  second  proglottis ;  the  pos- 
terior proglottis  shows  the  uterus 
partly  filled  with  eggs,  as  well 
as  the  cirrus  sac  and  the  vagina. 


'  In  Iceland  28  per  cent,  of  the  dogs  are  infected  with  this  Taenia,  in  Lyons  71  per  cent., 
in  Zurich  3*9  per  cent.,  in  Berlin  i  per  cent.,  and  in  Copenhagen  0*4  per  cent.  In  Australia 
even  40  to  50  per  cent,  of  the  dogs  are  affected.  It  is,  however,  a  question  whether,  in  addi- 
tion to  TcBnia  echinococcus,  a  second  analogous  form  is  not  involved,  as  the  form  from  Canis 
dingo  attains  a  length  of  10  to  30  mm. 


346 


THE   ANIMAL   PARASITES   OF   MAN 


man.  As  echinococcus  disease  in  man  is  always  very  dangerous,  it 
would  be  a  matter  of  general  interest  to  prevent  dogs  being  infected 
by  destroying  the  echinococci,^  and  all  measures  would  be  justifiable 
which  would  diminish  the  superfluous  number  of  house-dogs  (for 
instance,  high  taxes) ;  measures  should  also  be  adopted  to  limit  the 
association  of  men  with  dogs,  particularly  in  such  frequented  places 
as  restaurants,  railway  carriages  and  tram-cars. 

Echinococcus  is  very  common  in  slaughtered  animals  ;  in  Germany,  however, 
the  figures  in  the  reports  of  the  abattoirs  present  an  erroneous  view  in  so  far  as, 
besides  the  total  number  of  animals  slaughtered,  only  the  numbers  of  those  organs 
(liver  and  lungs)  are  published  that  were  so  severely  infected  with  echinococci  that, 
even  when  the  parasites  were  "  shelled  "  out,  the  flesh  could  not  be  placed  upon  the 
market  and  was  therefore  "condemned." 

In  Berlin  the  following  animals  were  slaughtered  : — 


Year 

1889—90 

1890—91 

1891—92 

1892-93 

1896-97 

1902 

Oxen  ... 
Sheep... 
Pigs    ... 

154,218 
430,362 
442,115 

124,593 
37i,Q43 
472,859 

136,368 
367,933 
530,551 

142,874 
355,949 
518,073 

146,612 
395,769 
694,170 

153,748 
434,155 
778,538 

During  the  same  years  the  following  were  condemned   in  consequence  of  being 
infected  with  echinococci  : — 


Lung 

Liver 

Lung 

Liver 

Lung 

Liver 

Lung 

Liver 

Lung 

Liver 

Lung 

Liver 

Oxen  ... 
Sheep... 
Pigs    ... 

7,266 

5,479 
6,523 

2,418 
2,742 
5,078 

5,792 

4,595 
5,083 

1,938 
2,059 

3,735 

4,497 
4,435 
6,037 

1,721 
1,669 
4,374 

2,563 
3,33^ 
6,785 

739 
1,161 
4,312 

3,284 
4,561 

7,888 

1,156 

1,939 
5,398 

2,507 
11,138 

9,544 

791 

4,437 
9,233 

Nevertheless  there  are  statistics  that  give  the  total  number  of  animals  infected 
with  echinococcus  : — 


Author 

Place 

Oxen 

Sheep 

Pigs 

Langrich    ... 

Olt...         

Steuding    ... 
Prettner      

Rostock  i.  M. 
Stettin 

Gotha 

Prague 

26- 2  per  cent. 

7-1 
24'6 
23-2 

37-0  per  cent. 

258 

35*4 

5"5 

5-4  per  cent. 
7-3 
21-4 

? 

In  Giistrow,  in  Mecklenburg,  half  of  the  animals  slaughtered  are  said  to  be 
infected  with  echinococcus;  in  Wismar  25  per  cent,  of  the  oxen,  15  per  cent,  of  the 
sheep  and  5  per  cent,  of  the  pigs  are  infected  ;  according  to  Mayer,  in  Leipzig, 
379  per  cent,  native  pigs,  24*47  per  cent.  Hungarian  pigs,  and  I3'09  per  cent,  of  sheep 
were  infected  with  echinococcus  ;  at  the  same  time  it  was  stated  that  in  regard  to 
the  native  pigs  the  liver  was  more  frequently  affected  than  the  lungs  (s'Si  per  cent. 
as  compared  with  0*26  per  cent.) ;  in  sheep  the  lungs  were  more  frequently  infected 


'  Mosler,  F.,  '*  Ueb.  Mittel  z.  Bekampfg.  endem.  vork.   Echinococcuskrank. , 
vud.  Zeit.,  1889,  No.  72. 


Deutsch. 


STRUCTURE   AND    DEVELOPMENT   OF    ECHINOCOCCUS 


347 


(1271  per  cent,  to  373  per  cent.)>  whereas  in  the  Hungarian  pigs  both  organs  were 
almost  equally  infected  (1478  per  cent,  to  I2'03  per  cent.). 

The  data  of  Lichtenheld,  in  Leipzig,  give  the  frequency  with  which  various 
organs  were  affected,  as  shown  in  the  following  table  : — 


Lungs  ... 

Liver    ... 

Spleen 

Heart  ... 

Kidneys 

Subperitoneal  tissue 


Cattle 


69*3  per  cent 
27-0       ,, 

2-2       „ 

075     .. 

075     .. 


Pigs 


i6*2  per  cent. 
74*2       ,, 
3*2      „ 

3'2         ,, 

3-2       „ 


21*4  percent, 
72-0       „ 

27       M 

1-3       M 

1-3      » 

i'3       >. 


Sheep 


52*2  per  cent. 
44*9       ,, 
2*9       », 


Horses 


.S'5  percent. 
94*5       .. 


Structure  and  Development  of  Echinococcus  (Hydatid). 

An  echinococcus  is  a  spherical  or  roundish  bladder  full  of  a 
watery  liquid,  which  originates  by  liquefaction  of  the  oncosphere,  and 
in  man  may  attain  the  size  of  a  child's  head,  but  remains  smaller  in 


Fig.  251. — Echinococcus  veterinorwn:  the  fibrous  sac  enclosing  the 
echinococcus  has  been  opened  and  laid  back  in  five  parts,  so  that  the 
surface  of  the  bladder  worm  may  be  seen,  with  the  brood  capsules,  visible 
to  the  naked  eye,  showing  through  it.     Natural  size.     (After  Leuckart.) 

cattle  (the  size  of  an  orange  or  apple).  The  thin  wall  of  the  bladder 
is  composed  of  an  external  laminated  cuticle  (ectocyst)  and  an  internal 
germinal    or    parenchymatous    layer    (endocyst).     The    latter    again 


348 


THE   ANIMAL   PARASITES   OF   MAN 


exhibits  two  layers  :  an  outer  layer  of  small  cells  that  are  less  sharply 
defined,  and  an  inner  layer  of  larger  cells.  It  contains,  in  addition, 
calcareous  corpuscles,  muscular  fibres  and  excretory  vessels.  It  is 
rich  in  glycogen. 


^^^^^^Ig       ^^Ml^^ 


Fig.  252. 

Figs.  252  and  252A. — Diagrams  of  mode  of  formation  of  brood  capsule  and  scolices. 
(i)  Wall  of  mother  cyst,  consisting  of  ectocyst  and  endocy.>,t  ;  (2)  theoretical  stage  of  invagination 
of  wall;  (3)  a  brood  capsule  with  the  layers  of  the  wall  in  the  reverse  position  to  that  in  the 
mother  cyst ;  {4)  evagination  of  wall ;  (5)  invagination  ;  (6)  fusion  to  form  the  solid  scolex  ; 
(7)  invagination  of  fore-part  of  scolex  into  hind-part.  [Note. — The  size  of  the  scolex  is  much 
out  of  proportion  to  the  brood  capsule.)     (Stephens.) 


STRUCTURE   AND   DEVELOPMENT   OF   ECHINOCOCCUS 


349 


The  development  in  cattle  often  remains  stationary  at  the  bladder 
stage,  and  they  are  then  called  "acephalocysts,"  or  Echinococciis  cysticus 
sterilis.  According  to  Lichtenheld,  sterile  cysts  occur  in  80  per  cent, 
of  cases  in  cattle,  in  20  per  cent,  in  pigs,  and  in  7-5  per  cent,  in  sheep. 
In  other  cases  large  numbers  of  small,  hollow  BROOD  capsules  are 
formed  in  the  germ  layer,  but  are  not  arranged  in  any  particular  order. 
The  order  of  the  layers  is  just  the  reverse  in  them  to  what  it  is  in  the 
parent  cyst,  that  is  to  say,  they  have  inside  a  thin  non-laminated  cuticle 
and  the  parenchymatous  layer  on  their  external  surface.  These, 
theoretically  at  least,  may  be  regarded  as  invaginations  of  the  bladder 
wall   giving    rise    to    a    cavity   with    the   cuticle    internal    and    the 


parenchymatous  layer  external.     If 
we   should    then    get    an    isolated 
cavity    with    cuticle    internal    and 
parenchymatous  layer  external,  as 
in    the    brood    capsule    (fig.   252). 


we  suppose  the  orifice  to  close, 


in 


ilMlill 


iJiiiiiiyiiiiii'iiii^iiiiiilB 


Fig.  252A. 

If  we  next  suppose  an  evagination  of  the  wall  of  the  brood  capsule  to 
occur  at  one  point  we  should  get  a  hollow  process  lined  with  cuticle  ; 
at  the  bottom  of  this  we  get  the  scolex  and  booklets  formed,  and  a 
little  higher  up  the  tube  the  suckers  (fig.  252,  4).  If  this  hollow  scolex 
is  now  pictured  as  being  invaginated  we  get  a  hollow  scolex  covered 
with  cuticle  and  lined  by  a  parenchymatous  layer  projecting  into  the 
cavity  of  the  brood  capsule.  The  two  sides  of  this  hollow  scolex  now 
fuse  and  we  get  a  solid  scolex  projecting  into  the  cavity.  Finally,  if 
we  imagine  once  more  the  rostellum  and  suckers  invaginated  into  the 
posterior  part  of  the  scolex  we  get  tlie  condition  as  frequently  found 
in  the  brood  capsules,/.^.,  a  scolex  covered  with  cuticle  projecting  into 
the  cavity,  with  the  rostellum  and  suckers  invaginated  into  the  posterior 
portion  of  the  scolex  (fig.  252A,  7). 


350 


THE   ANIMAL    PARASITES   OF   MAN 


A  large  hydatid  may  contain  many  thousands  of  brood  capsules. 
Each  brood  capsule  is  about  as  big  as  a  pin's  head,  and  may  contain 
ten  to  thirty  or  more  scolices.  The  delicate  wall  of  the  brood  capsules 
may  rupture,  so  that  the  scolices  are  now  free  in  the  mother  cyst.  These 
free  scolices  and  also  free  brood  capsules  constitute  what  is  known  as 
"  hydatid  sand,"  which  settles  at  the  bottom  of  a  glass  when  hydatid 
fluid  is  poured  into  it.  This  form  occurs  chiefly  in  domesticated 
animals  and  is  termed  E.  veierinortuu,    Rud.,  or  E.  cysticus  fertilis. 

In  man,  and  only  rarely  in  cattle,  the  mother  cyst  first  forms 
^'  daughter  cysts"  {E.  honiinis,  Rud.  [fig.  255]),  which,  though  smaller 
than  the  "  mother  cyst,"  re- 
semble it  in  the  structure  of 
their  walls  ;  thus  they  are 
covered  externally  by  a  lami- 
nated cuticle  and  internally 
by  the  parenchymatous  layer. 
They  originate  : 


Fig.  253. — Section  through 
an  invaginated  echinococcus 
scolex.  Cf.  fig.  252A,  7.  X  300. 
(After  Deve.) 


Fig.  254. — A  piece  of  the  wall  of  an  Echinococcus 
veterinorum  stretched  out  and  seen  from  the  internal 
surface.  A  few  brood  capsules  (the  outline  of  which 
is  only  faintly  shown),  with  scolices  directed  towards 
their  interior  and  exterior.     50/1. 


(i)  Between  the  laminae  of  the  cuticle  of  the  mother  cyst  from 
small,  detached  portions  of  the  parenchymatous  layer  ;  during  their 
growth  they  bulge  inwardly  or  outwardly  and  may  separate  them- 
selves entirely  from  their  parent  cyst.  In  the  latter  case  they  lie 
between  the  mother  cyst  and  the  capsule  of  connective  tissue 
formed  by  the  host  (E.  granulosus  or  E.  hydatidosiis  exogeniis) ; 
when  growing  inwardly  they  reach  the  interior  of  the  mother  cyst 
(£.  hydatidosiis  endogenus).  Their  number  is  very  variable  and  does 
not  depend  on  the  size  of  the  mother  cyst.  They  are  as  big  as,  or 
bigger  than,  gooseberries. 

(2)  According  to  some  authors,  endogenous  daughter  cysts  arise  also 


STRUCTURE   AND   DEVELOPMENT   OF   ECHINOCOCCUS 


351 


from  a  metamorphosis  of  scolices  that  have  separated  off  from  the  brood 
capsule.  This  takes  place  in  the  following  way  :  Fluid  accumulates 
in  the  interior  of  the  scolex,  so  that  eventually  nothing  remains  except 
a  sac  consisting  of  cuticle  lined  by  parenchyma.  The  cuticle  gradually 
thickens  and  several  layers  form  (tig.  257). 


Fig.  255* — Echinococcus  hoviinis  in  the  liver.  The  fibrous  capsule  and 
the  wall  of  the  echinococcus  have  been  incised,  so  that  the  endogenous 
daughter  cysts  may  be  seen.     Reduced.     (After  Oslertag,  from  Thoma.) 

(3)  Transformation  of  Brood  Capsules  into  Daughter  Cysts. — This  is 
also  held  to  be  possible  by  various  observers.  New  epithelial  layers  are 
deposited  between  the  cuticle 
which  lines  the  brood  capsule 
and  the  outer  parenchymatous 
layer.  This  parenchymatous 
layer  gradually  disappears  and  a 
newparenchymatous  layerforms 
in  the  interior  from  the  paren- 
chyma of  the  scolex  or  scolices. 
Although  it  appears  strange  that 
a  completely  formed  scolex  with 
specifically  differentiated  tissues 
and  organs  should  retrogress 
to  more  primitively  organized 
matter,  and  again  become  a 
proliferating  bladder,  yet  we  can 
hardly  doubt  that  the  older  observations,  regarding  such  a  vesicular 
metamorphosis,  of  Bremser  (1819),  v.  Siebold  (1837),  Naunyn  (1862), 
Rasmusser  (1866),  Leuckart  (1881),  Alexinsky  (1898),  Riemann  (1899), 
Deve  (1901),  and  Perroncito  (1902)  are  correct. 


Fig.  256. — Section  through  an  echinococcus 
scolex  in  process  of  vesicular  metamorphosis, 
twenty-six  days  after  insertion  in  the  pleural 
cavity,      x  250.     (After  Deve.) 


352 


THE   ANIMAL   PARASITES   OF  MAN 


(4)  Further,  a  fourth  method  of  formation  of  daughter  cysts  is 
described  by  Naunyn  as  occurring  in  sterile  hydatids,  i.e.,  those  con- 
taining no  brood  capsules.  In  this  case  a  portion  of  the  mother  wall 
of  the  hydatid  gets  constricted  off. 


Fig.  257. 

Ficis.  257  and  257A. — Diagram  of  transformation  of  a  scolex  into  a  daughter  cyst  (i  to  3)  : 
I,  scolex  in  brood  capsule  ;  2,  liquefaction  of  scolex  ;  3,  daughter  cyst ;  and  (4  to  6)  of  a  broou 
capsule  into  a  daughter  cyst :  4,  brood  capsule  with  scolex  ;  5,  deposition  of  new  epithelial 
layers  on  the  inner  layer  of  the  parenchyma  ;  b,  disappearance  of  outer  parenchyma  and  formation 
of  inner  parenchyma  from  the  parenchyma  of  scolex,  which  has  now  disappeared.  {Note. — 
The  scolices  are  out  of  proportion  to  the  brood  capsules  and  to  the  daughter  cysts.     (Stephens.) 

It  has  also  been  established  that  not  only  daughter  cysts  transplanted 
into  animals  develop  further  (Lebedeff,  Andrejew,  Stadnitzky,  Alexinsky, 
Riemann),  but  that  this  also  holds  good  if  only  hydatid  scolices  from 


STRUCTURE   AND   DEVELOPMENT   OF   ECHINOCOCCUS 


353 


man  or  animals  are  transplanted  into  animals  (rabbits).  They  develop 
into  echinococci  and  can  then  give  rise  to  brood  capsules  and  scolices. 
As  Deve  further  established,  hydatid  scolices  are  not  capable  of 
developing  in  guinea-pigs,  while  corresponding  experiments  with 
rabbits  are  in  the  large  majority  of  cases  successful  where  the  scolices 
are  introduced  subcutaneously  or  into  the  pleural  or  peritoneal  cavities. 
It  is  only  in  the  case  of  daughter  cysts  that  further  growth  is  obtained 
in  the  case  of  guinea-pigs.  Finally  it  appears,  as  has  been  already 
stated,  that  brood  capsules  can  transform  themselves  into  daughter 
cysts,  but  according  to  Deve  only  within  the  mother  cyst,  not  aftei 
transplantation.  Daughter  cysts  that  have  been  formed  in  the  mother 
cyst  of  man  and  animals  behave  themselves  just  as  the  mother  cyst 

does,  i.e.,  they  can  remain  sterile, 
r™  or  give  rise  to  brood  capsules 

l|lL         and    scolices,     or    even    again 
^        to    fresh    cysts — granddaughter 


Fig.  257A. 

cysts.  The  mother  cyst  can  also  die,  so  that  the  daughter  cysts  then 
lie  in  the  cavity  of  the  connective  tissue  capsule.  The  number  of  the 
daughter  cysts  in  either  case  may  attain  several  thousands. 

The  echinococcus  fluid,  which  originally  is  formed  from  the  blood  of  the  host,  is 
light  yellow,  with  a  neutral  or  slightly  acid  reaction  ;  its  specific  gravity  averages  1009 
to  1015.  It  contains  about  I'S  per  cent,  of  inorganic  salts,  half  of  which  is  common 
salt ;  in  addition  (besides  water)  it  contains  sugar,  inosite,  leucine,  tyrosin,  succinic 
acid  (associated  with  lime  or  soda)  and  albumens  which  are  not  coagulated  by  heat  ; 
occasionally  also  the  fluid  has  been  found  to  contain  haematoidin  and  uric  acid  salts 
(in  echinococcus  of  the  kidneys),  which  doubtless  demonstrates  that  the  echinococcus 
liciuid  originates  from  the  host.  It  has  been  generally  assumed  that  echinococcus 
fluid  contains  a  toxic  substance  the  escape  of  which  into  the  body  cavity  (at  operation 
or  by  bursting  of  a  hydatid  cyst)  produces  more  or  less  severe  symptoms  (fever, 
peritonitis,  urticaria),  so  much   so   that   one  speaks  of  hydatid  intoxication.     The 

23      . 


354 


THE   ANIMAL   PARASITES   OF    MAN 


investigations  of  Robert,  Joest,  etc.,  have,  however,  shown  the  harmlessness  of  fresh 
undecomposed  hydatid  and  cysticercus  fluid  for  rabbits,  mice  and  guinea-pigs,  whether 
inoculated  intraperitoneally,  subcutaneously  or  intravenously.  Contrary  data  or 
clinical  experience  must  accordingly  depend  on  other  factors. 

According  to  the  researches  of  Leuckart,  the  growth  of  the 
echinococcus  is  very  slow  ;  four  weeks  after  infection  the  average 
size  is  only  0*25  to  0-35  mm.,  at  the  age  of  eight  weeks  it  is  i  to 
2-5  mm.,  and  at  this  period  the  formation  of  the  central  cavity  com- 
mences ;  at  the  age  of  five  months,  and  with  a  size  of  15  to  20  mm., 
the  first  brood  capsules  with  scolices  are  formed.  The  conse- 
quence of  this  gradual  increase  of  size  is  that  the  organ  attacked 
can  maintain  its  functions  by  vicarious  hypertrophy,  and  that  many 
echinococci  induce  no  special  symptoms  and  cannot  even  be 
diagnosed,  the  latter  circumstance  being  due  to  their  hidden  position. 

The  echinococcus  cannot  be  said  to  be  scarce  in  man,  as  is 
shown  by  the  following  table  for  Central  Europe  : — 


Place 

Period 

No.  of 
post-mortems 

No.  of  cases 
of  echino. 

Percentage 

Rostock             

1861—83 

1,026 

25 

2-43 

Greifswald         

1862-93 

3.429 

51 

1-48 

Jena        

1866—87 

4,998 

42 

0-84 

Breslau 

1866—76 

5,128 

39 

0761 

Berlin 

1859-68 

4,770 

33 

0-69 

Wurzburg          

— 

2,280 

II 

0-48 

Gottingen          

— 

639 

3 

0-469 

Dresden 

1852—62 

1,939 

7 

036 

Munich 

1854-87 

I4,>83 

35 

0-2S 

Vienna 

i860 

1,229 

3 

024 

Prague 

— 

1,287 

3 

023 

Kiel        

1872—87 

3,581 

7 

0-19 

Ziirich,  Basle,  Berne 

— 

7,982 

II 

013 

Erlangen            

1862—73 

1,755 

2 

o-ii 

These,  however,  are  only  cases  that  have  become  known  by  post- 
mortem ;  in  addition,  there  are  cases  that  have  been  treated  medically, 
of  which  there  are  a  few  statements,  at  all  events  relating  to  the 
principal  districts  of  Germany.  According  to  Madelung,  one  case 
of  echinococcus  occurs  in  every  1,056  inhabitants  in  the  town  of 
Rostock,  in  the  district  of  Rostock  one  to  every  1,283,  in  Schwerin 
one  to  every  5,887,  and  in  Ludwigsort  one  to  every  23,685  ;  according 
to  Peiper,  in  Upper  Pomerania  one  case  occurs  to  every  3,336, 
in  the  district  of  Greifswald  one  to  every  1,535  inhabitants.  The 
northern  districts  of  Pomerania  are  more  affected  than  the  southern 
ones. 

Accordingly,  echinococcus  is  also  considerably  more  frequent  in  cattle  in 
Pomerania.  On  an  average  in  Germany  io"39  per  cent,  oxen,  9-83  per  cent,  sheep, 
and  6" 47  per  cent,  pigs  are  infected,  whereas  in  Upper  Pomerania  3773  per  cent. 


STRUCTURE   AND   DEVELOl^MENT   OF   ECHINOCOCCUS 


355 


oxen,  27-1  per  cent,  sheep,  and  I2'8  per  cent,  pigs  are  infected  ;  in  Greifswald,  indeed, 
64*58  per  cent,  oxen,  51 '02  per  cent,  sheep,  but  only  4*93  per  cent,  pigs  are  infected. 
In  accordance  with  these  figures  Tcc?iia  echmococcus  must  be  frequent  in  dogs  in 
Ponierania,  especially  in  Upper  Pomerania  ;  on  the  other  hand,  the  conjecture  that 
the  frequency  of  echinococcus  in  Mecklenburg  is  explained  by  the  occurrence  of 
TcB7ua  echinococcus  in  foxes  has  not  been  confirmed,  as  the  fox  does  not  harbour  this 
worm  in  Mecklenburg. 

Beyond  the  European  continent,  echinococcus  is  frequent  in 
the  inhabitants  of  Iceland,  Argentine,  Paraguay  and  Austraha.  In 
Iceland,  according  to  Finsen,  i  in  every  43  inhabitants  is  affected 
with  echinococcus  ;  according  to  Jonassen  the  proportion  is  i  to  63  ; 
this  is  due  to  the  habits  of  the  people  of  Iceland  or,  in  fact, 
to  the  frequency  of  Tcviiia  echino- 
cocciis  in  dogs,  and  the  prevalence 
of  the  hydatid  in  cattle.  In  cer- 
tain districts  of  Australia  it  is 
just  as  frequent.  In  Cape  Colony, 
Egypt  and  Algeria  echinococcus 
is  not  rare,  but  it  is  scarce  in 
America  and  in  Asia,  with  the 
exception  of  the  nomadic  tribes 
of  Lake  Baikal. 

Echinococcus  attacks  persons 
of  every  age,  though  it  is  rare  in 
children  up  to  10  years  of  age 
and  in  old  people.  It  occurs 
most  frequently  between  the  ages 
of  21  and  40  years.  According 
to  all  statistics  it  preponderates 
in  women  (about  two-thirds  of  the 
cases).  The  liver  is  its  favourite 
seat  (57-1  per  cent,  of  the  cases)  ; 
next  in  order  come  the  lungs 
(8  per  cent.),  kidneys  (6  per  cent.), 

cranial  cavity,  genitalia,  organs  of  circulation,  spleen  (3*8  per  cent.),  etc. 
As  a  rule  one  organ  only  is  invaded  ;  multiple  occurrence  may  originate 
from  one  infection,  or  eventually  from  a  later  infection  (?),  or  it  may 
come  to  pass  that  from  some  cause  (through  the  spontaneous  rupture  of 
an  echinococcus,  or  the  rupture  of  one  caused  by  an  injury  or  surgical 
operation)  daughter  cysts,  brood  capsules  or  scolices  escape  into  the 
abdominal  cavity,^  where  they  settle  or  become  transformed  and  go 


Fig.  258. — Hooklets  of  echinococcus.  a,  of 
Echinococcus  veterinorum  ;  b,  of  Tania  echino- 
coccus^ three  weeks  after  infection  ;  c,  of  the 
adult  Tcenia  echinococcus ;  d,  the  three  forms 
of  hooklets  outlined  one  within  the  other. 
600; I.     (After  Leuckart.) 


'  In  such  cases  the  toxic  effects  of  the  echinococcus  fluid  usually — if  not  always — manifest 
themselves.  Such  effects  are  manifested  by  severe  symptoms  of  poisoning  being  set  up,  by 
urticaria,  peritonitis,  and  ascites,  and  not  infrequently  they  cause  a  fatal  termination. 


356  THE   ANIMAL   PARASITES   OF   MAN 

on  growing.  In  the  distribution  of  this  secondary  echinococcus  the 
great  powers  of  motiHty  of  the  free  scoHces  must  be  taken  into 
account  (Sabrazes,  Muratet,  and  Husnot). 

Human  echinococci  may  also  die  at  various  stages  of  development,  become 
caseous  or  calcified,  or  may  be  absorbed,  the  cause  for  this  being  either  disease  of 
the  hydatid  itself  or  inflammation  of  its  connective  tissue  capsule;  the  discovery  of 
the  laminated  cuticle,  which  has  great  powers  of  resistance,  or  the  finding  of  the 
hooklets  of  the  scolices  is  sufficient  to  form  a  conclusion  as  to  the  nature  of  such 
formations. 

Siebold  (1853)  was  the  first  to  rear  Tcenia  echinococcus  in  the  dog 
by  feeding  it  with  the  echinococcus  of  cattle  and  especially  of  sheep. 
Kuchenmeister,  van  Beneden,  Leuckart,  Railliet  and  others  obtained 
similar  results,  and  Thomas,  Naunyn,  Krabbe  and  Finsen  succeeded 
in  rearing  T.  cc/iiiiococciis  in  dogs  from  the  bladder  worms  of  human 
beings  ;  these  grow  comparatively  slowly  (one  to  three  months^)  and 
only  during  the  process  of  growth  develop  their  hooklets  in  their 
definite  form  (fig.  258).  It  lies  in  the  nature  of  things  that  dogs, 
whether  experimentally  or  naturally  infected,  almost  always  harbour 
T.  echinococcus  in  large  quantities.  That  cats  exceptionally  harbour 
these  worms  has  been  already  mentioned  (Deve).  Finally,  Leuckart 
infected  young  pigs  by  feeding  them  with  mature  segments. 

Echinococcus  multilocularis  (alveolar  colloid). 

In  addition  to  the  form  of  echinococcus  already  described,  and 
which  is  also  frequently  termed  Echinococcus  nnilocnlaris,  there  is  a 
second  form  which  occurs  in  man  as  well  as  in  animals,  and  which 
is  termed  E.  ninUilocnlaris,  s.  alveolavis  (alveolar  colloid). 

It  was  originally  regarded  as  a  tumour  ;  its  animal  nature  was  first 
established  by  Zeller  and  R.  Virchow.  The  parasite,  which  varies 
in  size  from  that  of  a  fist  to  a  child's  head,  presents  a  collection  of 
numerous  cysts,  measuring  between  o'l  and  3  to  4  mm.  to  5  mm. 
in  diameter,  which  are  embedded  at  first  in  a  soft,  connective  tissue 
stroma  ;  the  cut  surface  has  therefore  a  honeycomb  appearance.  The 
cysts  are  surrounded  by  a  pellucid  and  laminated  cuticle,  and  each 
according  to  its  size  encloses  either  a  small-celled  tissue  or  a  cavity 
lined  by  a  parenchymatous  layer  ;  the  fluid  contained  in  such  a  cavity 
may  be  transparent,  or  is  rendered  opaque  by  globules  of  fat,  bile- 
pigment,  haematoidin  and  fat  crystals.  According  to  some  authors 
all  or  most  of  these  cysts  intercommunicate  ;  others  state  that  this  is 
the  case  at  least  as  regards  the  cuticle.     The  scolices  are  by  no  means 


'  According  to  Perroncito  the  scolices  had  not  formed  proglottids  nine  days  after  feeding, 
but  the  latter  were  present  twenty-four  days  after  feeding,  although  the  formation  of  eggs  had 
not  begun. 


ECHINOCOCCUS   MULTILOCULARIS  357 

found  in  all  the  cysts,  and  when  present  only  a  few,  rarely  half,  of  the 
cysts  contain  scolices  (one  or  more) ;  it  is  supposed  that  at  least  some 
of  these  scolices  are  formed  in  brood  capsules,  and  that  the  former 
are  capable  of  undergoing  a  cystic  metamorphosis. 

One  circumstance  is  peculiar  to  the  multilocular  echinococcus 
of  man,  namely,  the  disintegration  that  sets  in  at  certain  stages  ;  in 
the  centre  of  the  parasite  a  cavity  forms  that  frequently  becomes  very 
large  and  is  filled  with  a  purulent  or  brownish  or  brownish-green 
viscid  fluid  ;  in  this  fluid  one  finds  shreds  of  the  wall  of  the  cavity, 
calcareous  bodies,  echinococcus  cysts,  also  scolices  and  booklets,  as 
well  as  fat  globules  and  crystals  of  haematoidin,  margarine  and 
cholesterin  and  concretions  of  lime.  Such  ulcerative  processes, 
according  to  Ostertag,  are  never  present  in  the  multilocular  echino- 
coccus of  oxen,^  in  which  the  separate  cysts  are  larger  and  the 
connective  tissue  integument  less  powerfully  developed. 


Fig.  259. — Echinococcus  multilocularis  in  the  liver  of  the  ox.     Natural  size. 
(After  Ostertag.) 

Hardly  anything  positive  is  know^n  with  regard  to  the  development 
of  the  alveolar  echinococcus  ;  its  peculiar  conformation  is  attributed 
by  some  to  enormous  infection  of  oncospheres,  by  others  to  the 
abnormal  situation  of  one  oncosphere  ;  a  few  authors  ascribe  it  to 
infection  of  lymphatic  vessels,  others  to  infection  of  the  biliary  ducts 
or  to  peculiarities  of  the  surrounding  hepatic  tissue  ;  Leuckart  ascribes 
it  to  a  grape-like  variety  of  form  which  continues  budding  ;  a  few 
more  recent  authors  consider  multilocular  echinococcus  to  be  specific- 
ally different  from  unilocular  echinococcus,  and  therefore  also 
different  the  species  of  Taenia  arising  from  them.  Melnikow- 
Raswedenkow  is  also  of  this  opinion.  According  to  this  author  the 
oncospheres   infect   the    lumen    of   a  branch    of    the  portal  vein   in 

1  This  may  perhaps  be  explained  by  the  fact  that  the  hosts  are  slaughtered  before  the 
parasites  have  attained  the  size  or  other  conditions  necessary  to  disintegration. 


358  THE   ANIMAL   PARASITES   OF   MAN 

Glisson's  capsule  of  the  liver  and  grow  into  an  irregularly  sliaped 
formation  (chitinous  coil),  which  breaks  through  the  vascular  walls 
and  thus  forms  the  alveoli.  So  far  the  data  coincide  well  with 
Leuckart's  opinion  of  the  original  grape-like  form  of  the  Echiiiococcns 
multilocularis  ;  according  to  Melnikow-Raswedenkow  the  *'  granular 
protoplasmic  substance"  (parenchymatous  layer)  is  not  only  present 
in  the  interior  of  the  loculi  but  also  outside,  and,  moreover,  "ovoid 
embryos "  are  supposed  to  develop  in  the  chitinous  coils,  which, 
"  thanks  to  their  amoeboid  movements,  reach  the  lumen  of  a  vessel, 
where,  under  favourable  circumstances,  they  begin  to  develop 
further,"  that  is  to  say,  they  become  ''  chitinous  cysts  with  fantastic 
outlines,"  or  also  ^'  single-chambered  chitinous  cysts  "  ;  scolices  may 
develop  in  both.  Deve,  however,  considers  that  these  embryos  are 
only  prolongations  of  the  protoplasmic  layer  which  secondarily 
cuticularize. 

The  multilocular  echinococcus,  which  in  man  produces  a  severe  disease  and 
almost  always  leads  to  premature  death,  infects  most  frequently  the  liver,  but  may 
also  be  found  primarily  in  the  brain,  the  spleen  and  the  suprarenal  capsule  ;  from 
the  liver  by  means  of  metastasis  it  may  reach  the  most  various  organs,  especially 
those  of  the  abdomen,  but  also  the  lungs,  the  heart,  etc.  Up  to  1902,  235  cases  have 
been  described  and  up  to  1906,  265,  being  70  from  Russia,  56  from  Bavaria,  32  from 
Switzerland,  30  from  the  Austrian  Alps,  25  from  Wiirtemberg  ;  the  remaining  cases 
are  distributed  over  Central  Germany,  Baden,  Alsace,  France,  Upper  Italy,  North 
America.  In  some  the  origin  is  doubtful  ;  in  any  case  after  Russia,  the  mountainous 
South  of  Europe  is  the  principal  region  of  distribution.  As  to  the  domesticated 
animals,  the  same  parasite  is  found  principally  in  the  ox  (according  to  Meyer,  in 
Leipzig,  in  7  per  cent,  of  the  oxen  affected  with  echinococcus)  ;  it  is  rarer  in  the 
sheep  and  very  scarce  in  the  pig. 

It  has  already  been  mentioned  above  that  recently  the  multilocular 
echinococcus  has  been  stated  to  be  specifically  different  from  hydatid 
or  unilocular  echinococcus.  To  this  may  be  added  the  fact  that 
Mangold,  who  fed  a  young  pig  with  oncospheres  of  a  Taenia  reared 
from  the  multilocular  echinococcus,  found  two  growths  in  the  liver 
four  months  later,  which  he  took  to  be  E.  multilocularis,  and  con- 
sequently one  has  to  assume  the  existence  of  two  different  worms. 
The  chief  defender  of  this  view,  already  put  forward  by  Vogler, 
Mangold,  and  Miiller,  is  Possett.  He  bases  his  opinions  on  (i)  the 
more  restricted  distribution  of  the  multilocular  hydatid,  the  former 
occurring  in  districts  where  only  cattle  are  raised,  the  latter  where 
sheep-breeding  is  established  ;  (2)  that  those  engaged  in  looking  after 
sheep  are  attacked  by  multilocular,  whereas  those  looking  after  cattle 
are  attacked  by  unilocular  hydatid  ;  (3)  that  among  the  cases  of 
unilocular  hydatid  occurring  in  the  distribution  areas  of  multilocular 
hydatid  no  transitions  between  the  two  forms  are  observed  ;  (4)  on 
the  difference  in  the  hooks  both  in  the  hydatid  as-well  as  in  the  Taenia 


SERUM   DIAGNOSIS   OF   ECHINOCOCCUS  359 

stage  ;  the  hooks  of  Tcviiia  echinococcus  are  plump,  sharply  curved^ 
and  have  a  short  posterior  root  process  the  length  of  which  is  to  that 
of  the  total  length  as  i  to  47,  whereas  on  the  contrary  the  hooks  of  the 
alveolar  echinococcus  are  more  slender,  slightly  bent,  and  have  a 
long  posterior  root  process  (i  to  2-5)  ;  and  (5)  on  the  form  of  the 
uterus,  which  in  the  alveolar  Taenia  has  the  form  of  a  spherically 
distended  sac  anteriorly. 

Serum  Diagnosis   of   Echinococcus. 

(i)  Precipitin  Reaction. —M\x  equal  parts  of  hydatid  fluid  (of  the 
sheep)  and  serum  of  patient.  Keep  at  37°  C.  The  reaction  is  not 
decisive  as  it  may  be  given   by  normal  sera. 

(2)  Complement  Deviation. — Required  :  (i)  Hydatid  fluid  of  sheep 
(antigen),  (2)  guinea-pig  complement,  (3)  patient's  serum,  (4)  red 
cells  of  sheep,  (5)  haemolytic  serum  (of  rabbit)  against  sheep's  red 
cells,  (6)  o*8  per  cent,  salt  solution.  Mix  the  antigen  +  patient's 
serum  (heated)  +  complement  +  salt  solution  at  37°  C.  for  one  hour. 
Add  red  cells  of  sheep  +  haemolytic  serum.  Allow  to  stand  for 
half  an  hour  at  37°  C.  It  is  imperative  to  make  adequate  control 
observations.  An  example  will  indicate  the  method.  Salt  solution 
1*3  c.c.  4-  patient's  serum  (heated)  0*2  c.c.  +  hydatid  fluid  0*4  c.c.  + 
complement  ci  c.c.  of  serum  diluted  to  a  quarter  strength  +  haemo- 
lytic serum  and  red  cell  emulsion  i  c.c.  Result  :  no  haemolysis,  i.e.^ 
the  patient's  serum  contains  specific  (echinococcus)  antibodies. 


360  THE   ANIMAL   PARASITES   OF  MAN 


C.     NEMATHELMINTHES. 

J.    W.    W.    STEPHENS,    M.D.,    B.C.,    D.P.H. 

Bil.ATERALLY  Symmetrical  animals,  without  limbs  and  with  a  body  cavity  in 
which  the  gut  or  other  organs  float.     They  are  generally  cylindrical. 

Class.     NEMATODA. 
Nemathelminthes  with  an  alimentary  canal. 

Nematodes  are  as  a  rule  elongated  round  worms  of  a  filiform  or  fusiform  shape  ; 
their  length  varies  according  to  the  species  from  about  i  mm.  to  40  to  80  cm. 
The  outer  surface  of  the  body  is  smooth  or  annulated,  and  at  certain  points  provided 
with  papilla?,  occasionally  also  with  bristles  and  alar  appendages.  The  anterior  end 
■carrying  the  oral  aperture  is  usually  rather  slender,  occasionally  quite  thin  ;  the 
posterior  end  is  pointed  or  rounded  ;  the  anus,  as  a  rule,  lies  somewhat  in  front  of 
the  posterior  extremity.  The  sexes  are  almost  always  separate,  and  the  male  can 
<is  a  rule  be  easily  distinguished  from  the  female  because  the  former  is  smaller  and 
more  slender,  its  posterior  extremity  is  often  spiral  or  incurved,  or  carries  an  alar 
appendage,  whereas  the  female  is  larger  and  thicker,  and  its  posterior  extremity 
is  straight.  In  the  male  the  genitalia  open  into  the  anus  ;  the  sexual  orifice  of  the 
female  opens  ventrally  along  the  median  line  in  the  anterior  half  of  the  body,  in  the 
middle,  or  a  little  further  back.  Both  sexes,  moreover,  have  an  orifice,  the  excretory 
pore,  which  is  situated  ventrally  in  the  median  line  and  about  the  level  of  the 
<esophageal  nerve  ring. 

In  large  species,  even  with  the  naked  eye,  two  lighter  transparent  bands — the 
lateral  lines — may  be  distinguished  ;  they  run  along  the  sides  of  the  body  from  the 
anterior  to  the  posterior  end,  while  two  other  bands,  the  median  lines,  running 
along  the  ventral  and  dorsal  mid-lines,  are  less  evident  ;  in  exceptional  cases  there 
are  also  four  sub-median  lines.  These  bands  or  lines  are  inward  projections  of  the 
ectoderm,  and  in  them  lie  the  nerves  and  excretory  vessels  (fig.  260). 

Some  Nematodes  live  free  in  fresh  or  salt  water,  in  soil,  mud  or  decaying 
vegetable  matter,  others  parasitically  in  the  most  various  organs  of  animals, 
frequently  also  in  plants. 

Anatomy  of  the  Nematodes. 

All  the  Nematodes  are  covered  by  (i)  a  cuticle,  which  in  the 
small  species  is  thin  and  delicate,  while  in  the  larger  species  it  is 
thickened,  and  may  consist  of  several  layers  of  complicated  structure. 
Canalicular  pores  do  not  occur.  According  to  general  opinion, 
which  is  confirmed  by  the  history  of  development,  the  cuticle  is  a 
product  of  (2)  the  epithelium  or  ectoderm  that  had  formerly  existed 
or  is  still  found  beneath  it;  in  young  specimens  and  small  species  it 
IS  perceptible,  but  in  older  worms  it  frequently  alters  so  considerably 


ANATOMY  OF  THE  NEMATODES  36 1 

that  not  only  do  the  borders  of  the  cells  disappear/  but  a  fine  fibrous 
difterentiation  appears  in  their  cytoplasm.  The  matrix  or  ectoderm 
then  has  the  appearance  of  an  ectodermal  syncytium  permeated  by 
fibres  and  strewn  with  nuclei,  so  that  it  is  hardly  distinguishable  from 
the  tissue  of  (3)  the  cutis,  which  is  always  present,  though  developed 
to  a  varying  degree.  Both  layers,  matrix  and  cutis,  project  internally 
as  ridges  and  form  the  lateral  lines,  while  the  less  marked  median 
lines  are  produced  apparently  only  by  the  ectoderm  (fig.  260). 

Unicellular  cutaneous  glands  are  known  in  parasitic  as  well  as  in 
free-living  species  ;  they  vary  in  number  and  arrangement,  and  are 
found  discharging  some  at  the  anterior  extremity  and  others  in  the 
vicinity  of  the  genital  orifices.  In  other  cases  large  numbers  of  them 
are  present  along  the  lateral  lines ;  they  are  strongly  developed  in 
most  of  the  Trichotrachelidcv,  where  they  discharge  either  along  a 
part  of  the  ventral  surface  or  along  the  lateral  and  median  lines  ;  they 
are  placed  so  closely  together  that  the  ridges  of  the  cuticle  perforated 
by  the  orifices  have  long  been  known,  and  have  been  described,  as 
^^  rodlet  borders,"  or  *'  fields  of  rods." 

As  the  cutis  is  immediately  adjacent  to  (4)  the  dermo-muscular 
TUBE  the  simple  layer  of  the  muscular  cells  is  divided  into  four  quad- 
rants by  the  longitudinal  lines — two  dorsal  and  two  ventral  (fig.  260). 
The  muscles  are  in  the  simplest  cases  large  rhomboid  cells  that  lie 
two  by  two  in  each  quadrant,  so  that  on  transverse  section  of  the 
entire  worm  only  eight  cells  are  perceptible.  The  outer  border  of 
the  cells  is  converted  into  contractile  fibrils,  while  the  contiguous 
inner  portion  has  remained  protoplasmic,  and  contains  the  nucleus. 
In  large  species  the  muscular  cells  do  not  only  increase  in  length 
{up  to  3  mm.)  and  in  number  in  every  quadrant,  but  their  contractile 
portion  curves  up  to  form  a  groove  (like  that  of  a  dead  leaf)  thereby 
even  becoming  thicker;  simultaneously  space  is  gained  for  more 
cells,  the  protoplasmic  parts  of  these  cells  (on  transverse  section)  pro- 
ject out  of  the  grooves  like  vesicles.  In  all  cases  there  is  only  one 
layer  of  longitudinal  muscular  cells,  which, by  contracting,  can  shorten 
the  body  or,  by  contracting  one  side,  can  bend  it.  In  the  latter  case 
the  muscles  of  the  opposite  side  have  an  antagonistic  effect,  or  when 
all  the  muscles  are  contracted,  the  elasticity  of  the  cuticle  acts  in  the 
same  way.  Special  muscles  exist  at  the  beginning  of  the  gut  and  at 
sections  of  the  genital  apparatus. 

The  existence  of  a  cavity  between  the  body  and  the  gut  wall  has 
hitherto  been  generally  assumed,  and  has  been  referred  to  the  cleavage 
cavity,  and  consequently  designated  as  a  primary  body  cavity.     More 

'  In  the  Ascaridcc  isolated  epidermal  cells  grow  to  a  considerable  size,  and  have  to  do  with 
the  sensory  apparatus  of  the  lips  (Goldschmidt). 


36: 


THE    ANIMAT.    PARASITES    OF    MAN 


recent  investigators,  however,  state  that  such  a  cavity  does  not  exist, 
but  that  the  space  between  the  longitudinal  muscles  or  their  proto- 
plasmic portions  and  the  gut  epithelium  is  filled  by  a  complicated 
'isolation  tissue."  This  in  the  main  proceeds  from  a  large  cell  {Is., 
fig.  262)  which  lies  directly  behind  the  nerve  ring  dorsal  to  the 
oesophagus,  and  consists  of  a  system  of  lamellae  which  sheathe  the 
muscles  and  penetrate  through  them  to  the  cutis  and  also  cover  the 
gut  in  a  thin  layer. 

We  may  now  consider  the  ''tuft-like"  or  '' phagocytic "  organs, 
which  attain  i  cm.  in  size,  and  consist  of  four,  six,  or  even  more 
ramified  cells,  which   lie   close  to  the  walls   of   the   body  (fig.   261). 


Fig.  260. — Diagram  of  a  transverse  section  of  Asian's 
lumbricoides^  showing  thick  cuticle,  and  beneath  it  the 
matrix  or  syncytial  ectoderm.  The  flat  intestine  is  in 
the  middle,  and  to  the  right  and  left  near  it  in  the  body 
wall  the  lateral  lines  with  excretory  vessel  and  lateral 
nerves ;  above  and  below  in  the  centre  the  dorsal  or 
ventral  median  lines  with  the  nerves  radiating  to  the 
muscles,  also  the  muscle  cells  with  their  striated  outer 
contractile  portion  and  inner  nucleated  vesicular  proto- 
plasmic portion.     About  50/1.     (After  Brandes.) 


Fig.  261.  —  Anterior 
end  of  an  Ascaris  niegalo- 
cephala  cut  open  and 
showing  the  four  tuft- 
like organs  lying  on  the 
lateral  lines.  Natural  size. 
(After  Nassonow.) 


They  are  found  either  only  in  the  anterior  part  of  the  body  (Ascaris)^ 
or  throughout  the  whole  length  of  the  body  (Strongylus,  syn.,  Sclero- 
stomum),  and  their  position  usually  corresponds  to  the  lateral  lines. 
In  some  species  there  are  small  protoplasmic  cells  on  the  processes 
of  these  organs.  In  consequence  of  their  size  they  can  be  recognized 
with  the  naked  eye,  especially  when  they  are  loaded  with  granules  of 
stain  (carmine,  Indian  ink)  injected  into  the  body  cavity. 


ANATOMY   OF   THE   NEMATODES  363 

Intestinal  canal. — The  oral  aperture,  which  is  situated  at  the 
tip  of  the  anterior  extremity,  is  frequently  surrounded  by  thick  lips, 
or  small  bristles,  or  papillae  ;  it  leads  to  a  more  or  less  strongly 
developed  buccal  cavity,  which  is  lined  by  a  continuation  of  the  body 
cuticle,  and  which  in  some  species  is  provided  with  ''  teeth,"  repre- 
senting differentiated  portions  of  the  cuticle. 

The  cesophagus  (fig.  262),  which  arises  from  the  base  of  the  oral 
cavity,  is  as  a  rule  a  short,  bottle-shaped  tube  with  triradiate  lumen  ; 
its  wall  is  chiefiy  composed  of  radiating  muscular  fibres,  which  give 
it  the  appearance  of  being  transversely  striped  when  viewed  from  the 
surface.  There  exist  also  in  its  wall  three  large  gland  cells  (oeso- 
phageal glands)  and  nerves  arising  from  the  lateral  lines  and  running 
forward.  The  radial  fibres  cause  a  dilatation  of  the  lumen,  and 
exercise  an  effect  antagonistic  to  the  elasticity  of  the  cuticle  lining 
the  inner  surface.  The  latter  has  its  own  particular  layer,  which  is 
not  in  direct  connection  with  that  of  the  oral  cavity.  Special  dilator 
muscles,  arising  from  the  dermo-muscular  tube  and  situated  at  the 
commencement  of  the  oesophagus,  are  only  known  in  a  few  species. 
The  posterior  end  of  the  oesophagus  presents  a  bulb-like  dilatation, 
and  is  frequently  provided  with  small  chitinous  movable  valves.  In 
a  few  forms,  which  belong  to  the  Tricholrachelidcv  (Trichocephalus, 
Trichinella),  the  oesophagus  is  a  very  long  cuticular  tube,  beset  on  its 
dorsal  surface  with  a  series  of  large  nucleated  cells.  In  others  (Cucul- 
lanus,  Ascaris,  etc.),  a  tube,  the  so-called  glandular  stomach,  lined 
only  by  epithelial  cells,  follows  behind  the  muscular  oesophagus. 
This  glandular  stomach  is,  from  its  structure,  easily  distinguished 
from  the  mid-gut,  or  chyle  intestine,  which  is  likewise  cellular.  The 
so-called  mid-gut  is  a  tube  lined  by  flat,  cubical,  or  cylindrical  cells 
(fig.  260)  surrounded  by  *'  isolation  tissue  "  ;  its  transverse  section  is 
circular  or  flattened  dorso-ventrally  ;  the  lumen  may  run  in  a  straight 
line,  or  it  runs  a  sinuous  course  through  the  alternating  prominences 
of  the  then  flat  epithelial  cells. 

The  ectodermal  hind  gut  s,  as  a  rule,  very  short.  At  the  anal 
openmg  the  cuticle  and  the  subcuticular  layers  are  reflected  inwards, 
forming  the  lining  of  the  hind  gut.  In  large  species  the  subcuticular 
tissue  forms  large  cells  on  which  anteriorly  lie  in  addition  large 
''glandular  cells."  ^  n  the  male  the  ejaculatory  duct  opens  at  this 
point.  Around  the  end  part  of  the  gut,  either  on  the  chyle  intestine 
or  at  the  beginning  of  the  end  gut,  there  exists  a  sphincter  muscle 
arising  from  a  muscle  cell  which  acts  antagonistically  to  the  two 
diaphragm-like  dilator  muscle  cells  which  stretch  from  the  gut  to  the 

'  In  Ankylostomes  according  to  Looss  these  cells  have  no  glandular  function,  but  are 
ligaments. 


364 


THE   ANIMAL   PARASITES   OF   MAN 


Ml. 


body  wall.  In  many  species  large  stretches  of  tlie  gut  are  provided 
with  dilator  muscles.  There  is  sometimes  a  retrogressive  absorption 
of  the  gut  in  the  adult  stage  of  a  few  parasitic  species. 

Intestinal  c^ca  and  (esophageal  glands  sometimes  exist  as 

intestinal  appendages  ; 
the  former  are  tubular 
appendages  of  various 
size,  running  backwards 
or  forwards,  and  aris- 
ing from  the  posterior 
extremity  of  the  oeso- 
phagus. They  are  lack- 
ing in  many  species. 
The  oesophageal  glands 
are  unicellular  ;  a  dorsal 
and  two  sub  ventral 
glands  may  be  distin- 
guished according  to 
their  position  ;  as  a  rule 
they  open  into  the 
oesophagus  at  a  distance 
from  one  another.    The 

Fig.  262. — Transverse  &e.c\\OY\  ihxow^Ascajis  lumbticoides  1      j          r   j.i           1        a    y 

at  the  level   of  the  oesophagus  behind  the  nerve  ring.    Ctc,  uOCiy    Or    tne   glanO    lies 

cuticle;  Sc,  subcuticular  layer;  Ex.,  excretory  vessel ;  Is.,  in  the  bulb  of  the  OeSO- 

isolation  cell  and  the  system  of  lamellae  proceeding  from  it ;  ,                       •      4.1       j           1 

J/.,  muscles;  ML,  median  line  ;  SI.,  lateral  line.    Magnified.  Pl^^g'^S,  Or  m  tllC  dOrsal 

(After  Goidschmidt.)  ciil  -  ilc  -  sitc      arising 

from  it. 
The  nervous  system  is  sufficiently  known  in  a  few  species  only; 
it  consists  of  a  ring  containing  fifty  to  sixty  fibres  closely  surround- 
ing the  oesophagus,  various  groups  of  ganglion  cells,  and  a  certain 
number  of  nerves  extending  anteriorly  as  well  as  posteriorly.  The 
remarkably  small  number  of  fibres,  as  well  as  ganglion  cells,  is 
characteristic  of  the  nervous  system  of  all  Nematodes.  Immediately 
behind  the  oesophageal  ring  (fig.  263,  Lg.)  an  agglomeration  of  gan- 
glion cells  lies  at  either  side  (lateral  ganglia)  ;  part  of  their  off-shoots 
form  the  cesophageal  ring,  and  part  are  directed  posteriorly  and 
ventrally,  and  unite  partly  in  front  of  and  partly  at  the  back  of  the 
excretory  pore,  with  fibres  originating  direct  from  the  oesophageal 
ring,  and  passing  along  the  ventral  median  line  to  the  back  ;  these 
fibres  then  together  form  the  ventral  median  nerve  (fig.  263,  V. 111,11.). 
This  nerve,  originally  consisting  of  thirty  to  fifty  fibres,  becomes 
in  the  female  attenuated  quite  evenly  in  its  further  course.  There  is 
also  an  agglomeration  of  ganglion  cells  close  in  front  of  the  anus  (anal 
ganglia),  and  then-  the  median  nerve  divides  in  order  to  combine  with 


ANATOMY   OF   THE   NEMATODES 


365 


Lg.— 


Exp 


D.m.n. 


-  D.m. 


the  lateral  nerves  on  either  side.  In  the  male  the  median  nerve 
enlarges  to  nearly  the  original  number  of  fibres  in  front  of  the  anal 
ganglion,  which  contains  seven 
cells  ;  there  is  also  an  anal  ring 
embracing  the  terminal  gut,  and 
there  are  two  ganglion  cells  in  it  on 
each  side.  In  the  dorsal  median 
line  the  dorsal  median  nerve  is  alike 
in  both  sexes  ;  arising  in  front  with 
a  single  root  from  the  oesophageal 
ring,  it  gathers  its  fibres  from  the 
lateral  ganglia  ;  in  the  anterior  part 
of  the  body  it  consists  of  thirteen 
to  twenty  fibres  ;  in  the  posterior 
part  -of  the  body  the  fibres  are  re- 
duced to  four  or  six ;  behind  the 
anus  it  divides  and  combines  with 
the  lateral  nerves ;  the  latter 
consists  of  two  fascicles  at  either 
side  right  up  to  their  most  pos- 
terior extent — one  dorsal  and  one 
ventral — which  in  the  greater  part 
of  the  body  do  not  run  in,  but 
beside  the  lateral  lines,  and  exhibit 
a  different  origin  anteriorly.  The 
ventral  fascicle  at  each  side  branches 
off  from  the  ventral  median  nerve 
in  front  of  the  excretory  pore, 
whereas  the  dorsal  fascicles  origin- 
ate from  the  oesophageal  ring  close 
to  the  lateral  ganglia.  Each  of  the 
four  fascicles  contains  only  two  or 
three  fibres,  which  run  backwards 
parallel  to  the  lateral  lines;  a  few 
centimetres  in  front  of  the  caudal 
extremity  they  enter  the  lateral 
lines  and  remain  separate  from  one 
another  up  to  the  level  of  the  anal 
ganglion  ;  here  they  amalgamate  on 
either  side,  after  each  interpolating 
one  ganglion  cell,  with  the  single 
short  lateral  nerve  which  first  takes 

up  the  forked  ends  of  the  ventral  and  then  of  the  dorsal  median  nerve ; 
finally,  both  lateral  nerves  unite  wdth  each  other  at  the  back  in  an 
arch-like   manner. 


V.m  11. 


D.m. 


Fig.  263.— Schematic  representation  of 
the  nervous  system  of  a  male  Ascaris  mega- 
locephala.  A.,  anus;  Ag.^  anal  ganglion; 
C,  commissures  ;  D.m.n.,  dorsal  median 
nerve  ;  Exp.,  excretory  pore  ;  /V.,  oesopha- 
geal sensory  ring  ;  Lg. ,  lateral  gan}j;lia  ; 
Lit.,  lateral  nerve;  Sp.,  papilla;  V.m.n., 
ventral  median  nerve.     (After  Brandes.) 


366  THE   ANIMAL    PARASITES   OF   MAN 

In  the  male  each  ventral  part  of  the  lateral  nerves  becomes 
thickened  by  taking  up  fibres  from  the  ventral  nerves,  which  become 
thickened  posteriorly  to  the  nervus  bursalis,  which  towards  the 
middle  gives  off  a  mass  of  fibres  to  the  *^  genital  papillae  "  situated 
in  front  of  and  behind  the  anus  ;  the  number  of  these  fibres  averages 
eighty  to  100  ;  in  its  further  course  the  bursal  nerve  resembles  the 
corresponding  ventral  part  of  the  lateral  nerves  of  the  female. 

The  ventral  and  dorsal  nerves  are  connected  by  a  number  of 
semicircular  commissures,  which  originate  from  the  ventral  nerves 
and  serve  to  supply  the  dorsal  nerve,  which  is  always  being 
decreased  by  fibres  departing  from  it.  It  is  remarkable  that  these 
commissures  are  not  placed  symmetrically,  and  their  position  also 
is  different  in  the  two  sexes ;  in  the  female  there  are  thirty-one  on 
the  right  side  and  only  thirteen  on  the  left  side.  In  the  male  there 
are  thirty-three  commissures  on  the  right  side  and  fourteen  on  the 
left,  which  run  into  the  subcuticular  layer,  generally  in  pairs,  and 
usually  cross  at  the  level  of  the  lateral  lines. 

The  fibres  of  the  two  median  nerves  are  chiefly  motor ;  fascicular 
processes  run  from  each  protoplasmic  part  of  the  muscular  cells  to 
the  median  nerves  ;  from  these  they  take  up  bundles  of  primitive 
fibrils,  which  separate,  pass  through  the  protoplasmic  part  and  enter 
the  contractile  part  (fig.  260).  One  part  of  the  fibrils,  however, 
penetrates  beyond  the  muscles  into  the  subcuticular  layer,  where 
they  form  a  network,  probably  of  a  sensory  nature,  with  contiguous 
fibrils.  Nerves  directed  anteriorly  finally  originate  from  the 
oesophageal  ring;  they  consist  each  of  three  fibres,  carry  three 
ganglion  cells  at  their  point  of  origin,  and  enter  the  sensory  organs 
of  the  three  papillae  surrounding  the  oral  aperture.  Two  of  these 
little  trunks  lie  in  the  lateral  lines,  the  remaining  four  are  situated 
in  the  middle  of  the  four  quadrants  (Nn.  sub-mediani  anteriores). 

Parasitic  species  lack  higher  ORGANS  OF  sense  ;  free-living  worms 
occasionally  have  two  rust-red  eyes,  sometimes  with  lenses,  at  the 
anterior  part  of  the  body.  In  addition  to  the  above-mentioned 
sensory  papillae  surrounding  the  oral  aperture  and  the  genital  papilla 
of  the  male  at  the  end  of  the  body,  another  pair  exist  in  the  vicinity 
of  the  lateral  ganglia,  the  ''  cervical  papillae,"  and  two  dorsal  papilla 
in  the  central  region  of  the  body  and  two  lateral  ones  near  the  tip  of 
the  tail  {Ascaridce).  The  differences  in  the  distribution  and  number 
of  the  sensory  papillae  serve  for  characterizing  the  larger  and  smaller 
groups  of  Nematodes. 

The  excretory  organs  of  the  Nematodes  are  variable.  In  a 
great  many  cases  the  apparatus  is  symmetrical,  and  consists  of  a  vessel 
commencing  in  the  posterior  extremity  in  each  lateral  line  (fig.  260), 
and  passing  anteriorly.     In  the  vicinity  of  the  anterior  extremity  both 


ANATOMY   OF   THE   NEMATODES  367 

tubes  pass  out  of  the  lateral  lines,  bend  ventrally,  and,  in  the  median 
ventral  line,  unite  into  a  short  vesicle  formed  by  an  ectodermal  cell — 
the  cavity  of  which  is  lined  by  a  continuation  of  the  cuticle  of  the 
body — which  opens  into  the  excretory  pore  (fig.  263,  Exp.).  Asym- 
metry is  occasioned  through  the  excretory  duct  proceeding  from  the 
ventral  pore  to  the  lateral  line,  and  it  here  proceeds  as  (or  takes  up) 
the  left  excretory  canal,  which  anteriorly  is  a  broader  tube  and  runs 
along  the  left  lateral  line  ;  shortly  before  its  union  with  the  excretory 
duct  it  throws  out  a  branch  to  the  right  towards  the  lateral  line, 
which,  however,  always  remains  weak,  and  runs  posteriorly  in  the 
right  lateral  line  ;  a  few  smaller  branches  in  addition  spring  from 
the  left  main  stem.  In  other  species  the  right  branch  is  completely 
suppressed ;  the  entire  organ  thus  lies  in  the  left  lateral  line,  and 
consists  of  the  excretory  duct,  which  occasionally  opens  quite  in  front 
near  the  lips,  as  well  as  the  excretory  canal,  which  throws  out  a 
number  of  lateral  branches. 

This  excretory  vesicle  is  a  single  elongated  or  horse-shoe-shaped 
cell,  with  a  large  nucleus  and  an  intracellular  tubular  system,  which  is 
connected  with  the  excretory  duct  arising  from  the  excretory  pore  on 
the  outer  surface  (fig.  326).  The  so-called  ventral  gland  is  the  only  ex- 
cretory organ  of  marine  Nematodes,  and  probably  represents  a  primitive 
form.  Goldschmidt,  who  has  investigated  the  excretory  apparatus 
of  Ascaris  lumbricoldes,  considers  that  the  vessels  running  in  the  lateral 
lines  are  only  ducts  to  which  belong  a  glandular  system  hitherto 
overlooked  or  otherwise  interpreted.  This  system  also  lies  in  the 
lateral  lines,  and  takes  the  form  of  tw^o  glandular  tracts,  forming 
a  syncytial  tissue  in  which  lie  the  ducts,  one  dorsal,  one  ventral. 
In  parts  these  tracts  are  connected  by  commissures,  although  their 
junction  with  the  excretory  vessels  cannot  be  clearly  made  out.  These 
statements,  however,  require  confirmation.  The  author  has  further 
found  that  the  anterior  ends  of  the  lateral  canals,  directly  before  they 
bend  ventrally,  anastomose  with  one  another  and  give  off  anteriorly 
a  small  blind  process,  which  can  be  interpreted  as  a  rudiment 
of  a  canal  coming  from  the  head  end,  and  as  a  matter  of  fact, 
according  to  Golowin,  such  anterior  excretory  canals  exist  in  a 
number  of  genera. 

In  a  number  of  Nematodes  (Cheiracanthus,  Capillaria,  Tricho- 
cephalus,  Trichinella,  etc.),  however,  special  excretory  organs  are 
lacking  ;  possibly  the  cutaneous  glands,  which  are  in  these  species 
generally  powerfully  developed,  replace  these  organs. 

Sexual  organs. — With  the  exception  of  a  few  species,  the 
NejTiatodes  are  sexually  differentiated. 

(a)  Female  Sexual  Organs. — The  sexual  orifice  (vulva),  surrounded 
by  thick  labia,  is,  as  a  rule,  ventral  and  varies  in  position  from  near  the 


368 


THE   ANIMAL   PARASITES   OE   MAN 


head  to  near  the  anus.  It  leads  into  a  short  or  long  vagina  (ectodermic), 
bifurcating  into  the  two  uteri,  which  may  be  long  or  short  ;  the  long 
filiform  ovaries  are  continuations  of  them  (fig.  264).  Further  there  is 
often,  e.g.,  in  Ankylostoma,  a  differentiation  into  the  following  parts  : 


-at. 


Vul\/a- 


Ut. 


'Ov. 


Fig.  264.  —  Diagram  o  f 
female  genitalia.  Ov.^  ovary  (in 
part);  Ovd.^  oviduct;  Rec.  sent., 
seminal  receptacle;  Ui.,  uterus 
(in  part)  ;  Ovj.,  ovejector  ;  Vag., 
vagina. 


Fig.  264A. — Diagram  of  male  genitalia  of  a  strongy- 
lid.  Test.,  testis  (in  part) ;  S.  V.,  seminal  vesicle  ;  e.g., 
cement  gland  surrounding  ejaculatory  duct ;  j/.,  spi- 
cules; c/.,  cloaca;  gnd.,  gubernaculum  ;  p.p. a..,  pulvil- 
lus  postanalis  ;  g.c.^  genital  cone  ;  l.d.,  dorsal  lateral 
line ;  /.z'.,  ventral  lateral  line  (the  bursa  is  not  shown). 


(i)  Ovejector :  the  specialized  portion  of  the  uterus  before  it  joins  the 
vagina ;  there  may  be  a  separate  one  for  each  uterus,  or  a  common 
one  for  both  uteri.  (2)  Seininal  receptacle  :  at  the  other  extremity  of 
the  uterus.     (3)  Oviduct:  a  narrow  tube  connecting  the  ovary  with 


ANATOMY   OF  THE   NEMATODES  369 

the  uterus  proper.  (For  the  explanation  of  the  terms  convergent  and 
divergent  uteri  vide  footnote  p.  432.)  Uterus  and  ovaries,  which  arise 
in  the  first  place  from  a  single  cell,  lie  between  the  body  wall  and  the 
gut  and  are  surrounded  by  connective  tissue.  In  some  species  (for 
instance,  Trichinella)  the  ovary  is  single. 

At  the  blind  end  of  the  ovary  there  is  a  mass  of  protoplasm  with  numerous  nuclei 
that  multiply  continuously.  Gradually  the  nuclei  arrange  themselves  in  longitudinal 
rows  (fig.  265)  and  the  protoplasm  commences  to  leave  the  periphery  and  surround 
each  nucleus.  The  nearer  to  the  uterus  the  more  progressive  is  this  loosening 
process,  until  club-shaped  cells  each  containing  a  nucleus  are  developed.  The  most 
pointed  end  of  each,  however,  is  still  attached  to  an  axial  fibre  of  protoplasm,  the 
rhackisj  probably  this  has  some  connection  with  the  nutrition  of  the  ova.  Finally 
the  ova  fall  off  and  reach  the  uterus,  where  they  are  fertilized  and  enclosed  in  shells. 

(6)  Male  Sexual  Organs. — There  is  never  more  than  one  testis 
(fig.  266),  which  is  a  straight  or  sinuous  tube  of  the  same  construction 
as  an  ovary,  and  in  which  the  mother  cells  originate  in  the  same 
manner  as  the  ova.  In  the  same  way  as  the  ovary  passes  into  the 
uterus,  so  does  the  testis  pass  into  the  spermatic  duct  ;  the  latter  is 
often  divided  into  the  somewhat  dilated  seminal  vesicle  and  into 
the  muscular  ductus  ejaculatorius,  which,  running  ventral  to  the 
intestine  backward  (fig.  267),  finally  opens  into  the  cloaca.  In 
many  species,  e.g.y  A.  duodenale,  the  ejaculatory  duct  is  surrounded  for 
a  greater  or  less  portion  of  its  extent  by  the  cement  gland,  the  secretion 
of  which  (brownish  or  blackish  in  colour)  serves  for  copulation.  The 
ejaculatory  duct  of  the  large  Ascaridce  is  for  the  most  of  its  course 
surrounded  by  a  muscular  network  which  takes  its  origin  from  the 
two  dilator  cells  of  the  gut  (fig. 
268  F.).  The  spermatozoa  of  the 
Nematodes,  it  may  be  noted,  only 
attain  their  full  development  after 
the  sperm  mother  cells  have  been 
conveyed  by  copulation  into  the 

,      •       r    i.i_      r  1  -i.   1-  T  Fig.  265. — Transverse  section  throueh  the 

uteri   of    the   female  genitalia.      In        ovarian    tube   of  Belascaris  caii  of   the  cat 

their   form    (sheathleSS,   capable  of        ^^     various     levels.     To     dimonstrate     the 
,      •  ,  ,-        V   ,1  ,jx       r  development  (right   to  left)  of  the  ova  and 

amoeboid  motion)  they  differ  from      of  the  rhachis.    Magnified, 
those  of  most  other  animals. 

Spicules. — The  male  genital  apparatus  is  also  provided  with  one 
or  two  sacs,  situated  on  the  dorsal  side  of  the  intestine,  and  opening 
into  the  cloaca.  In  each  sac  there  is  a  chitinous  rod-like  body,  the 
spicule.  Further,  in  many  cases  there  exists,  more  or  less  fixed  in  the 
dorsal  wall  of  the  cloaca,  a  chitinous  structure,  the  accessory  piece  or 
gubernaculiimf  the  latter  name  implying  its  function  of  guiding  the 
spicules  during  copulation  (fig.  264A).  A  special  muscular  apparatus, 
24 


370 


THE   ANIMAL   PARASITEvS   OF   MAN 


consisting  of  protractors  and  retractors,  moves  the  spicules.  The 
protractors  or  exsertors  in  the  large  Ascaridae  consist  of  four  flat 
band-like  muscles  which  surround  the  spicule  sac.  Two  long  muscle 
cells  which  arise  proportionally  far  forward  on  the  dorsal  side  of  the 
lateral  line  and  are  inserted  into  the  base  of  the  spicules    serve    as 

retractors.     The  spicules   can  be   projected 
Q  from  the  cloacal  orifice  (anus)  during  copu- 

lation, and  when  they  are  introduced  into 
the  vagina  they  serve  as  prehensile  organs, 
perhaps  also  as  stimulatory  organs. 


.sp. 

A. 
P. 

Fig.  266. —  Male  of  the 
rhabditic  form  of  An- 
giostovmm  nigrovenosuj/i. 
A.,  anus;  /.,  mid-gut; 
T.,  testicular  tube;  a, 
oral  orifice  ;  P.,  papillae  ; 
5^.,  spicule.  Magnified. 


Fig.  267. — Transverse  section  through  the 
posterior  extremity  of  the  body  of  Ascaris 
Imnbricoides  (male).  The  intestine  is  in  the 
middle,  and  the  lateral  lines  are  subjoined 
thereto ;  above  the  intestine  the  two  spicule 
sacs  are  seen  ;  below  is  the  ductus  ejaculatorius. 
The  muscular  fibres  are  between  the  lateral 
and  median  lines.     Magnified. 


Bursa  copulatrix. — The  males  in  many  genera  possess  epidermal 
wing-like  appendages  at  their  posterior  extremity.  These  are  supported 
by  elongated  tactile  papillae  called  ribs.  In  the  most  highly  developed 
bursae,  e.g.^  in  the  Strongylidce,  the  ribs  are  called  rays,  as  they  consist 
not  only  of  nerve  fibres  but  mainly  of  "  pulp,"  i.e.,  prolongations  of 
the  subcuticular  layer.  Bursae  are  either  open,  i.e.,  bilaterally 
symmetrical,  or  closed,  when  the  posterior  border  is  continuous  all 
round.  A  pseudo-bursa  is  one  unsupported  by  ribs  or  rays,  e.g.,  in 
Trichuris.  The  bursa  serves  as  an  organ  of  prehension  during 
copulation.  Some  forms,  moreover,  carry  a  sucker  at  the  posterior 
extremity  (e.g.,  Heterakis)  ;  in  others  the  spicules  and  other  pre- 
hensile organs  are  absent  ;  they  are  then  replaced  by  an  evertible 
cloaca,  e.g.,  Trichinella. 


DEVELOPMENT  OF  THE  NEMATODES 


371 


Development  op^  the   Nematodes. 

After  impregnation,  the  ovum  develops  around  itself  a  delicate 
membrane  (vitelline  membrane),  and  subsequently  an  egg-shell  is 
formed.  This  is  derived  either  as  a  secretion  from  the  uterine  wall 
or  it  is  a  further  differentiation  of  the  vitelline  membrane,  the  origi- 
nal single  membrane  splitting  into  two,  the  outer  becoming  the  egg- 
shell. Further  the  uterus  often  secretes  a  special  albuminous  covering 
around  the  egg-shell.  The  "yolk  "  granules  of  the  ovum  are  secre- 
tions of  the  protoplasm  of  the  ovum  itself  and  first  appear  when  the 
rhachis  is  formed.     In  certain  cases  ova  lie  in  follicles  or  capsules 


sp. 


SI. 


D. 


DiL 


Fig.  268. — Hind  end  of  a  male  Ascaris  lumbricoides  cut  across  at  the  level  of  the  dilator 
cells  of  the  gut.  Z>.,  gut;  Dil.^  dilator  cells  of  the  gut;  F.,  a  process  of  the  dilator  cells 
forming  a  network  over  the  vas  deferens;  SL,  lateral  line;  Sp.,  spicule;  Fa^.,  vas  deferens. 
The  anterior  end  of  the  worm  lies  to  the  right.     Magnified.     (After  Goldschmidt.) 


formed  of    epithelium  cells  derived  from  the  ovarian  tubes.     These 
cells  subsequently  fuse  and  form  a  membrane — the  CHORION. 

The  shape  of  the  completed  eggs  is  characteristic  of  the  different 
species,  and  therefore  a  single  egg  often  suffices  to  diagnose  the 
species.  According  to  the  species,  the  eggs  may  be  deposited  sooner 
or  later,  either  before  or  during  segmentation,  or  with  the  embryo 
perfectly  developed.  Only  a  few  species  are  viviparous,  e.g.^  Dracun- 
ciilus  medinensis,  Trichinella  spiralis;  in  the  other  Nematodes  the 
further  development  of  the  extruded  eggs  takes  place  after  various 


372  THE   ANIMAL   PARASITES   OF   MAN 

lengths  of  time  in  the  open,  in  moist  earth,  or  in  water.  Thick-shelled 
eggs  can  maintain  their  developmental  capacity  for  a  long  time,  even 
after  prolonged  desiccation. 

Finally,  a  nematode-like  embryo  develops,  which  usually  lies 
somewhat  coiled  up  within  the  shell,  and  varies  in  its  further 
development  according  to  the  species  to  which  it  belongs. 

In  the  simplest  forms,  as  in  the  free-living  Nematodes,  the 
embryos,  apart  from  their  size,  resemble  their  parents,  and  grow  up 
into  these  after  leaving  the  egg-shell.  In  many  parasitical  Nema- 
todes, however,  the  young  must  be  called  larvce,  as  they  present 
characters  which  are  subsequently  lost. 

The  manner  of  conveyance  of  the  eggs  or  the  embryos  contained 
in  them  after  they  have  left  the  body  into  the  definite  host  is  very 
different  in  the  various  species. 

(i)  Without  Intermediate  Host. — (a)  In  many  the  conveyance  into 
the  definite  host  is  effected  directly  after  the  larvae  have  developed 
within  the  eggs  ;  thus,  for  instance,  the  feeding  of  suitable  animals 
with  the  embryo-containing  eggs  of  species  of  Trichocephalus  and 
Ascaris  leads  to  an  infection  of  the  gut,  for  the  young  Trichocephali 
or  Ascarides  only  leave  the  egg-shell  when  they  have  attained  the 
intestine  of  the  final  host,  in  which  they  become  adult. 

In  other  cases  (6)  Ancylostoma,  Necator,  the  larvae  hatch  in  the 
open,  and  live  for  a  time  free,  changing  their  form  ;  they  grow,  cast 
their  skin,  and  finally  gain  the  intestine  of  the  host  by  means  of  water 
or  through  the  skin,  when  they  lose  their  larval  characters  and 
assume  the  structure  of  the  adult  worm. 

(c)  In  a  number  of  Nematodes,  however,  heterogony  occurs. 
This  terms  signifies  a  mode  of  development  in  which  two  structurally 
different  sexual  generations  of  the  same  species  alternate  with  each 
other.  To  these  appertains,  for  instance,  Angiostomum  (syn.  : 
Rhabdonema)  nigrovenosumy  which  lives  in  the  lungs  of  frogs  and 
toads  ;  this  Nematode  measures  about  i  cm.  in  length  and  is 
hermaphrodite  (protanrdic).  The  eggs  are  deposited  in  the  pulmonary 
cavity,  and  through  the  cilia  of  the  same  reach  the  oral  cavity,  where 
they  are  swallowed  and  thus  conveyed  into  the  intestine.  They  pass 
through  the  entire  gut,  and  are  finally  evacuated  with  the  faeces  ;  often, 
indeed,  the  young  themselves  emerge  from  the  egg-shell  within  the 
hind-gut  of  the  frogs.  These  young  forms  become  sexually  differ- 
entiated, remain  much  smaller  than  the  parent,  their  oesophagus  is 
differently  constructed  (rhabditis  form),  and  they  are  non-parasitic 
(fig.  266).  After  having  grown  in  the  open  they  copulate ;  the  males 
die  soon  after  copulation,  and  the  females  in  their  own  bodies  develop 
a  few  young,  which,  given  the  opportunity  to  get  into  frogs,  infect 
them,  and  are   transformed  into  the  hermaphroditic  Angiostomum. 


DEVELOPMENT   OF   THE   NEMATODES 


373 


The  same  manner  of  development  occurs  in  other  species  of  the  same 
genus,  and  also  in  the  case  of  Stroitgy hides  stercoralis. 

(2)  With  Ifiteiinediatc  Host. 
— {a)  Frequently,  however,  the 
larvae  of  Nematodes  make  use 
of  one  or  even  two  intermediate 
hosts  ;  their  condition  then 
resembles  that  of  Cestodes  or 
Trematodes,  excepting  that 
there  is  never  a  multiplication 
within  the  intermediate  hosts. 
The  larvae  become  encapsuled 
amongst  the  tissues  of  the  in- 
termediate host,  and  wait  till 
they  are  introduced  with  the 
latter  into  the  final  host.  For 
instance,  OlUdanns  tricuspis,  the 
adult  form  of  which  is  found  in 
cats,  previously  lives  encysted 
in  the  muscular  system  of  mice. 
Cucidlaniis  elegans,  which  at- 
tains the  adult  stage  in  fishes 
(perch,  etc.),  is  found  encysted 
in    species  of  Cyclops.     Other 

examples    of    species  that  require  an  intermediate  host    are    Filaria 
barter  of ti  and    Dractmculus   medinensis. 

Peculiar  conditions  prevail  in  the  case  of  {h)  Trichinella  spiralis. 
This  species,  which  in  its  adult  state  lives  in  the  intestine  of  man  and 
of  various  mammals,  is  viviparous  ;  the  young  Trichinae,  however,  do 
not  leave  the  intestine,  but  reach  the  intestinal  w^all  (Cerfontaine, 
Askanazy)  in  the  following  way  :  the  female  intestinal  Trichinae  bore 
into  the  intestinal  wall,  where  they  are  found  in  the  submucosa,  or 
in  the  lumen  of  the  dilated  lacteal  vessels.  Here  the  young  are  born, 
in  the  intestinal  wall,  and  leave  this  position  with  the  lymph  stream. 
Some  of  them,  no  doubt,  actively  bore  through  the  intestinal  wall, 
reaching  the  lymph  or  blood-stream,  or  even  pass  into  the  body 
cavity.  What  occurs  during  their  further  migrations  is  difficult  to 
say  at  present.  It  has  hitherto  been  maintained  that  the  wandering 
is  entirely  active ;  for  instance,  the  ligaturing  of  an  artery  would  be 
no  protection  against  the  part  of  the  body  supplied  by  such  artery 
being  invaded  by  Trichinella.  This  observation  cannot  be  otherwise 
explained  than  by  the  active  progress  of  the  young  Trichinella.  The 
question,  however,  may  be  mooted  as  to  where  and  when  the  worms 
quit  the  blood-vessels,  which  they  for  the  most  part  reach  through 


Fig.  269. — A  piece  of  the  trunk  muscle  of  the 
pig  with  encapsuled  embryonic  Trichinse.  Mag- 
nified. 


374  THE  ANIMAL   PARASITES   OF   MAN 

the  thoracic  duct,  the  natural  connection  between  tlie  vascular  system 
and  the  lymphatic  system,  to  wander  further  independently,  and 
ultimately  reach  the  muscular  system,  in  which  they  become  encysted 
(fig.  269).  Thus  the  progeny  does  not  leave  the  body  of  the  host 
inhabited  by  the  parents,  as  is  generally  the  case  amongst  helminthes, 
but  uses  it  as  an  intermediate  carrier  to  reach  another  host,  which 
is  then  the  final  host.  The  latter  may  belong  to  another  species, 
or  may  be  another  individual  of  the  same  species.  This  second 
migration    is,  of   course,  purely  passive. 

Classification  of  the  Nematoda. 

The  Nematodes  are  usually  divided  into  a  number  of  families,  some  of  which 
it  is  at  present  impossible  to  define  accurately  ;  moreover,  the  definition  of  many 
genera  is  also  in  an  unsatisfactory  state. 

Family.     Anguillulidae,  Gervais  and  van  Beneden,  1859. 

A  "family"  name  not  definable.  They  comprise  a  vast  number  of  small  forms, 
most  of  which  live  free  in  fresh  water,  in  soil,  or  in  macerating  substances  ;  amongst 
them  there  are  some  which  live  parasitically  on  plants,  more  rarely  on  animals. 
They  do  not  exceed  8  mm.  in  length.  The  large  majority  are  only  i  to  2  mm.,  or 
even  0*5  mm.  The  uterus  is  straight.  Eggs  in  the  uterus  at  one  time,  one  to 
four.  Genera  very  numerous,  but  many  of  them  insufficiently  defined  (Anguillula, 
Anguillulina,  Rhabditis,  Heterodera,  etc.). 

Family.     Angiostomidae,  Braun,  1895. 

Small  Nematodes  undefined  morphologically,  but  characterized  by  heterogony, 
i.e.,  there  is  a  free-living  "  rhabditic  "  generation  and  a  parasitic  "  filariform  "  genera- 
tion which  succeed  one  another  {e.g.,  Angiostomum,  Strongyloides,  Probstmayria). 

Family.     Gnathostomidae. 

Cuticle  covered  totally  or  partly  with  cuticular  laminae  fringed  posteriorly  with 
multiple  points.  Head  subglobular,  covered  with  simple  spines.  Two  spicules. 
Vulva  behind  middle  of  body,  parasitic  in  vertebrates,  especially  mammals  {e.g., 
Gnathostoma,  Tanqua,  Rictularia). 

Family.     Dracunculidae,  Leiper,  191 2. 

Males  very  small  in  proportion  to  females.  Anus  absent.  Vulva  absent  (.?). 
Genera  :    Dracunculus,  Icthyonema  (in  body  cavity  of  eel  and  other  fish). 

Family.     Filariidse,  Glaus,  1885. 

Long  thread-like  Nematodes.  Anus  present.  CEsophagus  without  bulb. 
Vulva  usually  in  anterior  half  of  body.     Two   ovaries.      Generally   ovoviviparous. 


CLASSIFICATION   OF   THE   NEMATODA  375 

Development  often  requires  an  intermediate  host.  This  family  is  at  present 
ill-defined,  but  has  been  already  subdivided  into  several  sub-families,  Filariince^ 
OnckocercincE,  Arduennince. 


Family.     Trichinellidae,  Stiles  and  Crane,  1910. 

CEsophagus  consisting  of  a  chain  of  single  cells,  the  lumen  of  the  fjesophagus 
passing  through  their  centre.  Ovary  single.  Vulva  at  junction  of  anterior  and 
posterior  portions.     Sub-families:    (i)   Trichurince^  (2)   Trichinellincc. 


Family.     Dioctophymldae. 

Body  anteriorly  armed  with  spines  or  unarmed  ;  mouth  without  lips,  with 
six,  twelve,  or  eighteen  papillae  in  one  or  two  circles ;  oesophagus  very  long 
without  a  bulb  ;  anus  terminal  in  female  ;  one  ovary  ;  vagina  very  long ;  spicule 
in  male  very  long ;  bursa  cup-shaped  without  rays  (Dioctophyme,  Hystrichis, 
Eustrongylides). 

Family.     Strongylidae,  Cobbold,  1864. 
Bursa,  supported  by  rays,  always  present.     Oviparous. 


Family.     Physalopteridae. 

Mouth    with   two   large   lips.      Bursa   with   supporting   papillae   in   form   of  a 
lanceolate  cuticular  expansion,  with  genus  Physaloptera. 


Family.     Ascaridae,  Cobbold,  1864. 

Rather   thick   Nematodes.      Mouth    with   three   lips — one   dorsal,   two   latere- 
ventral.     Sub-families  :  (i)  Ascarince,  (2)  Heterakince^  etc. 


Family.     Oxyuridae. 

Smallish  forms,  4  to  45  mm.,  with  cuticle  thickened  on  each  side  for  the 
whole  length  of  body  in  the  form  of  a  lateral  flange  or  wing.  CEsophagus  long 
with  a  well-marked  bulb  containing  a  valvular  apparatus.  Tail  end  of  female 
drawn  out  into  a  long  point.  Eggs  asymmetrical.  Males  very  sniall  (about 
2  mm.).     One  spicule.      Genera  :  Oxyuris,  Passalurus,  Ozolaimus,  Atractis,  etc. 

Mermithidcs,  greatly  elongated  "  Nematodes,"  which,  in  the  larval  stage,  are 
parasitic  in  insects,  but  in  their  adult  condition  are  free  living.  Cuticle  with 
diagonal  striation.  Without  an  open  mouth  or  anus.  Oral  papillae  present. 
Characteristic  eggs  with  two  processes,  ending  in  a  tuft  of  filaments.  Larvas 
with  a  movable  boring  spine  at  the  head  end, 

Gordiidce. — Long,  thread-like  "  Nematodes."  Mouth  and  anterior  portion  of 
gut  atrophied  in  adult.      Oral  papillae  absent. 


376 


THE   ANIMAL   PARASITES   OF   MAN 


The  Nematodes  Observed  in   Man. 


Family                     Sub-family 

Genus 

Species 

Anguillulida-     ...                 — 

Rhabditis 

R.pellio. 
R.  niellyi. 
Rhabditis  sp. 

Anguillula 

A.  aceti. 

Anguillulina 

A.  putrefaciens. 

AngiostotnidcB  ...                — 

Strongyloides 

St.  stercoralis. 

Gnathostomidce .»,                — 

Gnathostoma 

On.  siajnense. 
On.  spinigerum. 

Dracunculidd   ...                — 

Dracunculus 

D.  7nedinensis. 

FilariidcB           ...     Filar iince   ... 

Filaria       

F.  bancrofti. 
F.  deinarquayi. 
F.  taniguchi. 
F.  (?)  conjunctives. 

Group.     Agamofilaria    ... 

Ag.  georgiana. 
Ag.  palpebralis. 
Ag.  oculi  huniani. 
Ag.  labialis. 
F.     (?)   romanorum- 

orienialis. 
F.  (?)  kilimarce. 
F{l)sp.l 

{Mikrqfilaria) 

Mf.  powelli. 

Mf.  philippinensis. 

Setaria 

S.  equina. 

Loa 

L.  loa. 

A  canthocheilonema 

Ac.  Persians. 

Dirofilaria 

Di.  7nagalhdesi. 

Onchocercince 

Onchocerca 

0.  volvulus. 

Trichinellida     ...     Trichurince 

Trichuris 

T.  trichiura. 

TrichinellincE 

Trichinella... 

T.  spiralis. 

DioctophymidcE ...                 — 

Dioctophyme 

D.  gigas. 

Strongylidce       ...     Metastrongy lines  ... 

Metastf  -ongylus     . . . 

M.  apri. 

Trichostrongylince 

Trichostrongylus  ... 

T.  instabilis. 
T.  probolurus. 
T.  vitrinus. 

Hcemonchus 

H.  contortus. 

Mecistocirrus     {Ne- 

M.  fordi. 

matodirus) 

AncylostomincE 

Group.     CEsophagostomecE 

Ternidens  ... 

T.  deminutus. 

CEsophagostojnuJTt. . . 

CE.  brujnpti. 

GE.  stephanostomum 

var.  thomasi. 
CE.  apiostomum. 

Group.     Ancylostomece  ... 

Ancylostoma 

A.  duodenale. 
A.  ceylanicum. 
A.  braziliense. 

Group.     Bunostomece 

Necator 

N.  ainericanus. 
N.  exilidens. 

Group.     SyngamecE 

Synganius  ... 

S.  kingi. 

ANGUILLULID^ 


377 


P'amily 
Physalopterido' . . 

Ascaridcg 


Oxyurida 
Mermithidce 


Sub-family  Genus 

—  ...     Physaloptera 

Ascarincp    ...         ...     As  carts 


Toxascaris 
Belascaris  ... 

Lagocheilascaris 
Oxyuris 
Mermis 
{Agamomermis) 


Species 
P.  caucasica. 
P.  mordens. 
A.  lumbricoides. 
A.sp. 
A.  texana. 

A,  maritima. 
T.  limbata, 

B.  cati. 

B.  marginata. 
L.  minor, 
O.  vermicularis. 
M.  hominis  oris. 
A£:  restiformis. 


Family.     Anguillulidae. 

Genus.     Rhabditis,  Dujardin,   1845. 

Buccal  cavity  elongated,  with  lips.     Its  chitinous  wall  uniformly  thick.     Lateral 
lines  absent.     Males  with  bursa. 


Rhabditis  peliio,  Schneider,  1866. 

Syn. :    Pelodera  peliio,  Schn.,  1866;   Rhabditis  genitalis,  Scheiber,  1880; 
Rhabditis  peliio,  Schn.,  1866. 

Males  0*8  to  1*05  mm.  in  length  ;  females,  0*9  to  1*3  mm.  in  length. 
The  posterior  extremity  of  the  body  of  the  male  has  a  heart-shaped 
bursa,  and  seven  to  ten  ribs  on  each  side  ;  the  bursa  may,  however, 
be  lacking.  The  spicules  measure  o'027  to  0*033  ^^V^-  ii^  length,  but 
are  never  quite  alike.  The  posterior  extremity  of  the  female  is  long 
and  pointed  ;  the  vulva  lies  somewhat  behind  the  middle  of  the  body, 
the  ovary  is  single,  the  eggs  are  oval,  60  jm  by  35  fju. 

This  species  was  found  in  Stuhlweissenburg  by  Scheiber  in  the  acid  urine 
(containing  albumin,  pus  and  blood)  of  a  woman  suffering  from  pyelonephritis, 
pneumonia  and  acute  intestinal  catarrh  ;  the  observer  was  able  to  convince  himself 
that  the  Nematodes  which  were  found  during  the  whole  period  of  the  illness  lived 
in  the  vagina,  and  were  evacuated  with  the  urine. 

Oerley  proved  that  this  species  had  long  been  known  ;  during 
its  larval  stage  {Anguilhila  mucronata,  Grube,  1849)  it  lives  in  earth- 
worms;  in  its  adult  stage  it  lives  in  decomposing  matter  in  the  soil. 
By  introducing  individuals  of  this  species  into  the  vagina  of  mice, 
Oerley  succeeded  in  obtaining  infection  and  multiplication  (facultative 
parasitism).  These  Nematodes  must  in  some  such  way  have  got  into 
the  vagina  of  Scheiber's  patient. 

Two  other  cases  described  by  Baginsky  and  Peiper  probably 
belonged  to  the  same  or  a  nearly  related  species. 


378  THE   ANIMAL   PARASITES   OF   MAN 

Rhabditis  nieliyi,  Blanchard,  1885. 

Syn.  :  Leptodera  nieliyi,  Blanchard,  1885. 

In  1882  Nielly  had  a  cabin-boy,  aged  14,  under  observation 
in  Brest.  The  lad  had  never  left  the  neighbourhood  of  Brest, 
and  had  suffered  from  itching  papules  on  the  skin  for  five  or 
six  weeks  ;  in  the  papules  the  observer  found  one  or  several  rhab- 
dites,  measuring  0-33  mm.  in  length  by  0*30  mm.  in  breadth. 
Their  cuticle  presented  a  delicate  transverse  striation  ;  the  intestine 
was  the  only  internal  organ  recognizable,  and  it  opened  somewhat 
in  front  of  the  posterior  extremity.  Therefore,  it  must  have  belonged 
to  the  rhabditis-like  larva  of  a  Nematode,  the  adult  stage  of  which 
is  unknown. 

The  manner  of  infection  was  established  almost  certainly  by  a  further  obser- 
vation of  Nielly's  :  at  the  commencement  of  the  illness  small  Nematodes  were 
found  in  the  blood  of  the  patient ;  later  on,  however,  they  disappeared,  neither  were 
Nematodes  found  in  the  faeces,  urine  or  sputum.  Therefore  it  must  be  concluded 
that  the  cabin-boy,  who  was  in  the  habit  of  drinking  water  from  brooks,  had  thus 
ingested  embryo-containing  eggs  of  a  Nematode  ;  the  young  hatched  out  in  the 
intestine,  perforated  it,  reached  the  blood  and  then  settled  in  the  skin  ;  but,  on  the 
other  hand,  the  entry  may  have  been  direct  through  the  skin. 

In  connection  with  the  foregoing,  reference  should  be  made  to  a  communication 
by  Whittles,  insufficient  from  a  zoological  point  of  view.  In  a  case  of  hypertrophic 
gingivitis  occurring  in  a  female  patient,  aged  19,  who  had  never  left  Birmingham, 
he  found  Nematode  larvae  in  the  periosteum  of  the  upper  jaw,  which  was  excised 
after  extraction  of  the  right  premolar  ;  the  genital  rudiment  could  be  recognized  in 
them.  Similar  larvae  were  found  in  the  same  patient  in  abscesses  in  various  regions 
of  the  skin,  and  in  the  case  of  her  mother  in  the  blood.  The  author  considers  that 
the  infection  took  place  through  a  dog,  and  refers  to  the  case  of  O'Neil  (1875),  ^^^ 
found  Filariae  in  the  skin  (in  the  condition  known  as  "craw-craw"),  referred  by 
Manson  to  Filaria  Persians.  O'Neil's  case  was  quoted,  and  attributed  to  Filaria 
sanguinis  hominis.  In  conclusion,  the  author  states  that  he  has  repeatedly  found 
Nematode  larvae  in  the  blood  of  persons  who  suffered  from  pruritus;  in  his  opinion 
the  parasite  had  been  imported  through  the  agency  of  troops  returned  from  South 
Africa.  Glatzel  found  true  Filaria  larvae  in  a  pustule  of  a  cutaneous  eruption  of  the 
trunk  and  extremities  in  a  patient  at  Dar-es-Salam. 

Skin  diseases  which  are  caused  by  young  Nematodes  are  also  observed  in  dogs 
(Siedamgrotzky,  MoUer,  J.  G.  Schneider,  Kiinnemann),  foxes  (Leuckart),  and  horses 
(Semmer).  Ziirn  found  young  Nematodes  {AnguillulidcB)  also  in  pig's  flesh.  In 
Kiinnemann's  case  it  was  shown  that  the  adult  Rhabdites  lived  in  the  straw  upon 
which  the  dog  lay. 

Rhabditis,  sp. 

In  the  fluid  obtained  by  lavage  from  the  stomach  of  a  female  patient,  aged  16, 
suffering  from  ozaena,  O.  Frese  found  during  two  consecutive  months  Rhabdites  of 
various  ages,  0*275  to  0-64  mm.  in  length,  the  adults  all  with  eggs  ;  males  were 
not  found;  transmission  into  rabbit's  stomach  failed,  but  they  could  be  kept  alive  in 
much  diluted  hydrochloric  acid  (2  :  1,000)  for  several  weeks.     Neither  eggs  nor  larvae 


ANGUILLULA 


379 


appeared  in  the  faeces  of  the  patient.     The  nature  of  the  infection,  which  was  perhaps 
of  unique  occurrence,  remained  doubtful. 

Genus.     Anguillula,  Ehrenberg,  1826. 

Buccal  cavity  very  small,  without  lips.  Males  without  bursa,  but  with  a  series  of 
papillae.     Lateral  lines  absent. 

Anguillula  aceti,   Miiller,   1783. 

CuticLile  unstriped,  body  cylindrical,  anterior  end  tapering  but  little, 
posterior  end  long,  pointed.  Male  up  to  1*45  mm.  long,  0*024  to 
0*028  mm.  wide;  two  pre-anal  papillae,  one  post-anal ;  spicules  equal, 
curved,  0*038  mm.  long  ;  gubernaculum  present  ;  testis  extending  in 
front  of  mid-line  of  body.  Female  up  to  2*4  mm.  long,  0*040  to 
0*072  mm.  wide  ;  anterior  uterus  reaching  to  near  the  oesophagus, 
posterior  to  hind  gut.  Viviparous ;  embryos  in  both  or  only  in  one 
uterine  horn,  o'22  mm.  long,  0*012  mm.  broad. 

The  species  is  a  frequent  inhabitant  of  vinegar  (prepared  by  older  methods),  and 
was  once  observed  for  some  time  by  Stiles  and  Frankland  in  the  urine  of  a  woman  ; 
the  urine  had  an  acid  reaction,  and  once  had  a  distinct  odour  of  vinegar.  It  was 
assumed  that  the  patient,  who  was  hysterical  and  suffered  from  chronic  nephritis, 
employed  vaginal  douches  with  diluted  vinegar,  perhaps  to  deceive  her  physician  or 
to  protect  herself  against  conception.  According  to  Ward,  Billings  and  Miller  are 
said  to  have  reported  on  two  other  cases.  Ill-effects  which  might  be  connected  with 
the  vinegar  eel  {Anguillula  aceti)  were  not  present. 

Genus.     Anguillulina,  Gervais  and  Beneden,  1859. 

Syn.  :   Tylenchus^  Bastian,  1864. 

Characterized  by  the  possession  in  the  buccal  cavity  of  a  spine  knobbed  pos- 
teriorly ;  bursa  present  ;  uterus  asymmetrical.  Numerous  species  parasitic  in 
plants. 

Anguillulina  putrefaciens,  Kiihn,  1879. 

Syn.:    Tylenchtis  putrefaciens^  YjSSvi\\   Trichina  contorta^  Botkin,  1883. 

In  1883  Botkin  {Pet.klin.  Wochensckr.,  1883)  found  a  small  Nematode,  which  was, 
however,  entirely  mistaken,  in  the  material  vomited  by  a  Russian  ;  this  was  not  a 
species  of  Trichinella,  but  an  Anguillulina  living  in  onions  which  had  already,  in 
1879,  been  described  by  Kiihn  as  Tylenchus  putrefaciens ;  the  Nematodes  got  into 
the  stomach  with  the  onions,  causing  nausea  and  vomiting. 

Family.     Angiostomidae,  Braun,  1895. 
Genus.     Strongyloides,  Grassi,  1879. 

Syn.  :  Pseudorhabditis^  Perroncito,  1881  ;  Rhabdonema^  Leuckart,  1882,  p.p. 

The  genus  is  insufficiently  defined.  The  parasitic  form  possesses  a  simple  mouth 
opening  directly  into  the  long  cylindrical  oesophagus  which  occupies  the  anterior 
third  of  the  body.  The  free-living  forms  possess  a  small  buccal  cavity ;  the 
oesophagus  is  short,  with  a  double  bulb,  in  the  hinder  one  there  is  a  Y-shaped 
chitinous  valve  ;  two  spicules  of  equal  size. 


38o 


THE   ANIMAL   PARASITES   OF   MAN 


Syn. 


Strongyloides  stercoralis,  Bavay,  ^877. 

Anguillula  intestinalis  et  stercoralis,  Bavay,  1877  ;  Leptodera  intestinalis 


et   stercoralis,   Cobb. ; 
strongyloides,  Leuckart 


Pseudorhabditis  stercoralis,    Perroncito,   1881  ;   Rhabdonema 
1883  ;   Strongyloides  intestinalis,  Grassi,  1883  ;  Rhabdonema 
intestinale,  Blanchard,  1886. 

In  1876,  a  number  of  French 
soldiers  returned  to  Toulon  from 
Cochin  China  suffering  from 
severe  diarrhoea.  Dr.  Normand, 
under  whose  treatment  they 
were,  discovered  a  large  number 
of  Nematodes  in  the  evacuated 
faeces,     and     Bavay     described 


CEsophagus 
Intestine 


/ 


Excretory  pore 


Ovary,  anterior 


m 


Testis 


Ovary,  anterior 


Ovary,  anterior 
Ovary,  posterior 


,'»« 


Testis 


Anus 
Intes- 
tine 


.   Vesicula  seminalis 


Bursal  muscles 
Anal  papilla 


;''  / .   Papills 


.cU/- 


Spicule 

Ovary,  posterior       ^,-^«Hrw  .^-^i^aE^ 

^^^^^"^^^  Ovary,  posterior 

Fig.    271. — Strongyloides    ster- 

Fig.    270. — Strongyloides  stercoralis^   female:    parasitic     (:^ra//j,  male :  free-living  generation. 

generation  from  gut  of  man.      x  70.     (After  Looss.)         X   170.     (After  Looss.) 


them  as  Anguillula  stercoralis.  Soon  after  Normand,  at  the  post-?nortem  of  five 
patients  who  had  died  of  Cochin  China  diarrhcea.  found  numerous  other 
Nematodes  in  the  intestine,  from  the  stomach  to  the  rectum,  in  the  bile-ducts  and 
in  the  pancreas,  and  these  he  handed  over  to  Bavay.     The  latter  diagnosed  another 


STRONGYLOIDES   STERCORALIS  38 1 

species,  and  described  them  as  A.  intestinalis.  Both  forms  were  then  regarded 
as  the  cause  of  Cochin  China  diarrhoea  until,  in  1882,  Leuckart  was  able  to 
demonstrate  that  the  two  forms  are  only  two  succeeding  generations  of  the  same 
species,  of  which  the  one  {A.  intestinalis)  lives  parasiiically  in  the  intestine, 
whereas  its  young  {A.  stercoralis)  attain  the  open,  where  they  come  to  maturity 
and  propagate.  The  young  of  these  again  live  parasitically.  There  thus  exists 
the  same  heterogony  as  was  discovered  by  Leuckart  in  Angiostomutn  nigro- 
venosum  of  frogs,  which  heterogony,  indeed,  according  to  v.  Linstow,  appertains 
to  the  entile  family  of  the  Angiostotnidce. 

(i)  The  parasitic  generation  (strongyloid  or  filariform  ?)  is  quite 
colourless  and  cannot  be  seen  m  situ  even  with  a  lens.  To  detect 
them  it  is  necessary  to  scrape  the  mucosa  of  the  jejunum  and  examine 
the  scrapings  microscopically.  It  measures  2*2  mm.  in  length,  and 
34  yu,  to  70  yLt  in  breadth  ;  the  cuticle  is  finely  transversely  striated  ; 
the  mouth  is  surrounded  by  four  lips  ;  the  oesophagus  is  almost  cyl- 
indrical and  a  third  the  length  of  the  entire  body.  The  anus  opens 
shortly  in  front  of  the  pointed  posterior  extremity ;  the  vulva  is 
situated  at  junction  of  middle  and  posterior  thirds  of  the  body;  the 
uterus  has  no  special  ovejector ;  the  eggs  measure  50  //,  to  58  yu,  in 
length,  and  30  /i-  to  34  yu,  in  breadth,  and  lie  in  a  chain  one  behind 
the  other  (fig.  270).  As  in  the  case  of  Angiostomum  nigrovenosiim^ 
Leuckart  considers  this  stage  to  be  hermaphroditic,  the  testes 
degenerating  after  having  functioned;  other  authors  (Rovelli)  regard 
it  as  a  female  reproducing  by  parthenogenesis. 

(2)  The  free-living  generation  ( ^  and  ?  )  has  a  smooth  body, 
cylindrical,  somewhat  more  slender  at  the  anterior  extremity  and 
pointed  at  the  tail  end.  The  mouth  has  four  indistinct  lips ;  the 
oesophagus  is  short  with  a  double  (rhabditis-like)  bulb ;  there  is  a 
Y-shaped  valve  in  the  posterior  bulb  ;  the  anus  opens  in  front  of  the 
tail  end.  The  males  measure  07  mm.  in  length,  0*035  i^^n^-  ^^  breadth. 
Their  posterior  end  is  rolled  up  ;  the  two  brown  spicules  are  small 
(38  ft)  and  much  curved.  There  is  also  a  gubernaculum.  The 
females  measure  i  mm.  in  length  or  a  little  over ;  0*05  mm.  in  breadth. 
The  tail  end  is  straight  and  pointed  ;  the  vulva  lies  somewhat  behind 
the  middle  of  the  body.  The  yellowish,  thin-shelled  ova  measure 
70  fi  in  length  and  45  ^  in  breadth. 

As  Askanazy  has  shown,  the  parasitic  form  bores  deeply  into  the 
mucous  membrane  of  the  intestine,  and  frequently  into  the  epithelium 
of  Lieberkiihn's  glands,  both  for  nourishment  and  oviposition.  The 
eggs  then  develop  in  the  intestinal  wall.  The  eggs  which  are  found  in 
scrapings  from  the  mucosa  occur,  at  least  in  the  case  of  Strongyloides 
of  the  sheep,  in  chains  enclosed  in  a  thin  tube  or  sheath,  the  origin 
of  which  is  doubtful ;  possibly  it  is  the  uterus.  The  eggs  them- 
selves are  only  rarely  found  in  stools,  e.g.^  after  a  strong  purge.  The 
larvae,  which  are  hatched  out,  and  measure  0*2  to  0*25  mm.  long  by 


382 


THE   ANIMAL   PARASITES   OF   MAN 


o*oi6  mm.  broad,  again  reach  the  lumen  of  the  intestine,^  and  grow  to 
double  or  three  times  that  size,  until  they  are  passed  out  with  the 
faeces.  They  already  differ  from  the  parent  (  $  )  in  the  shape  (rhab- 
ditiform)   of   the   oesophagus.      When    the    external    temperature    is 

sufficiently  high   (26^  to 


:M 


Nerve-ring 


Excretoiy  pore 


Ovary,  anterior 


Ovary,  anterior 


Ovary,  posterior 


Ovary,  anterior 

Anterior  oviduct 

Anterior  receptaculum 
seminis 


Posterior  senainal  receptacle 


M'1 


)\aiy  posterior 


Fig.  272. — Strongyloides  stercoralis,  female  ;  free-living 
generation,      x   170.     (After  Looss.) 


35°  C),  they  become 
sexually  mature  after 
moulting.  In  about 
thirty  hours  they  are  com- 
pletely  developed  and 
copulate,  now  forming 
the  free-living  rhabditi- 
form  generation.  At 
lower  temperatures  the 
larvae  only  moult,  but 
do  not  escape  from  the 
old  cuticle   and  do    not 


\ 


Genital 
rudiment 


.\'-V\ 


\ 


Fig.  273. — Strongyloides 
stercoralis :  larva  from  fresh 
human  faeces,  x  310.  (After 
Looss.) 


develop  further.     At  a  temperature  of  about  25°  C.  only  some  of  the 
larvae  attain  maturity. 

'  As  a  case  published  by  Teissier  shows,  they  may  also  abnormally  appear  in  the  blood 
(Arch.  mtd.  exptr.  et  d'Aji.  path.,  1895,  vii,  p.  675). 


STRONGYLOIDES   STERCORALIS 


383 


The  females  of  the  free-Hving 
generation  (rhabditiform)  deposit  from 
thirty  to  forty  eggs,  which  develop 
rapidly,  sometimes  even  within  the 
uterus  in  the  case  of  old  females. 
After  the  larvae  have  emerged  from  the 
egg-shell,  they  measure  0*22  mm.  in 
length,  and  possess  the  characteristics 
of  the  parents  (rhabditiform  larvae). 
When  they  have  grown  to  0*55  mm. 
they  moult,  and  while  losing  their 
own  characteristics  they  acquire  the 
characteristics  of  their  parasitic  grand- 
parents (strongyloid  or  filariform). 
After  about  eight  days  the  free-living 
adult  generation  in  the  cultures  have 
disappeared,  and  all  the  rhabditiform 
larvae  have  been  transformed  into 
strongyloid  or  filariform  larvae  ;  they 
then  die  off  unless  they  reach  the 
intestine. 

This  cycle  of  development  holds 
good  for  Strongyloides  stercoralis  of 
tropical  origin  (Bavay,  Leuckart,  Leich- 
tenstern,  Zinn).  In  the  European 
Strongyloides  the  free-living  genera- 
tion, as  a  rule,  is  absent  (Grassi, 
Sonsino,  Leichtenstern,  Braun)  ;  the 
rhabditis-like  larvae  evacuated  with  the 
faeces  are  transformed  into  the  strongy- 
loid or  filariform  type  of  larva  (in 
cultures  which  are  easily  made)  which 
will  only  become  adult  if  introduced 
into  man. 

So  that  we  have  these  two  cycles  : 
{A)  (i)  $  parasitic,  (2)  eggs,  the  rhab- 
ditiform larvae  in  faeces,  (3)  free- 
living   strongyloid  or  filariform  larva, 

(4)  $  parasitic.  (B)  (i)  (2)  (3)  as  before, 
then    (4)   adult    $    and    ^ ,   free  living, 

(5)  eggs,      (6)      rhabditiform     larva, 

(7)  strongyloid    or    filariform     larva, 

(8)  ?    parasitic. 

Infection  of  man   results  not  only 


Nerve-ring 


Subventral  oeso- 
phageal glands 

Excretory  pore 


CEsophagus 


--  Genital  rudiment 


Anus 


Fig.  274. — Strongyloides  stercoralis: 
mature  filariform  larva  showing  long 
transparent  oesophagus,  slender  granu- 
lar intestine  and  characteristic  tip  to 
the  tail  ending  in  two  small  points. 
X  620.     (After  Looss). 


384  THE   ANIMAL   PARASITES   OF   MAN 

from  direct  entry  into  the  stomach  but  also,  according  to  van  Durme 
and  Looss,  through  the  skin. 

Occurrence  i?i  Man.—k.'s,  already  mentioned,  Strongy hides  stercoralis  was 
first  observed  in  persons  suffering  from  so-called  Cochin  China  diarrhoea. 
From  the  enormous  numbers  of  parasites  evacuated  with  the  faeces,  the  cause 
of  the  disease  was  apparently  evident.  It  appeared,  however,  that  only  some  of 
the  soldiers  returning  from  Cochin  China  and  Martinique,  and  suffering  from 
diarrhoea,  harboured  Strongyloides  (Chauvin).  Breton  made  the  same  observations 
in  Cochin  China  and  found  that  only  io'4  per  cent,  of  cases  of  chronic  dysentery, 
and  8*8  per  cent,  of  chronic  diarrhoea,  show  Strongyloides.  Normand,  moreover, 
found  that  only  a  few  of  the  Europeans  residing  in  Cochin  China  are  exempt 
from  S.  intestinalis,  yet  the  people  exhibit  no  intestinal  symptoms  ;  if,  however, 
from  any  cause  a  catarrhal  condition  of  the  intestine  supervenes  the  condition 
is  changed,  the  parasites  appear  in  larger  numbers,  and  the  disorder  is 
considerably   intensified. 

S.  intesiinalis,  besides  being  present  in  the  Indo-China  region,  also  occurs 
in  the  Antilles,  in  Brazil,  Africa,  and  Europe  ;  in  1878  it  was  discovered 
in  Italy  by  Grassi  and  C.  and  E.  Parona ;  in  1880  it  was  also  found  in  the 
labourers  working  at  the  St.  Gothard  tunnel.  It  was  imported  into  Germany, 
Belgium,  and  the  Netherlands  by  Italian  labourers.  One  sporadic  case  has  been 
observed  in  East  Prussia,  and  the  worm  has  also  been  reported  from  Siberia. 

In  mammals  the  following  species  are  found  :  Probstmayria  {Strongyloides) 
vivipara,  Ransom,  1907,  in  Equus  caballus ;  Strongyloides  fulleborni,  \.  Linst.,  in 
Anthropopithecus  troglodytes  and  Cynocephalus  babuin. 

Their  development  is,  so  far  as  is  known,  the  same  as  that  of  Strongyloides 
stercoralis  (v.  Linstow,  Centralbl.  f.  Bakt.^  Path.  u.  Infektionsk..,  19055  Orig.  xxxviii, 
P-  532). 

Family.     Gnathostonriidae. 
Genus.     Gnathostonna,  Owen,  1836. 

Syn,  :  Ckeiracanthus,  Diesing,  1839. 

Easily  recognizable  by  the  numerous  spines  which  cover  the  entire  body  or 
only  the  anterior  extremity,  and  terminate  in  several  points  ;  head  globular  and 
beset  with  bristles  ;  mouth  with  two  lips  ;  two  spicules  ;  vulva  situated  behind  the 
middle  of  the  body. 

Gnathostoma  siamense,  Levinsen,  1889. 

Syn.  :  Ckeiracanthus  siamense^  Lev.,  1889. 

Female  measures  9  mm.  in  length,  i  mm.  in  breadth.  There  are 
eight  rows  of  simple  spines  on  the  head ;  the  armature  of  spines  extends 
over  the  anterior  third  of  the  body  only  ;  each  spine  on  the  anterior 
region  of  the  body  spreads  into  three  points,  of  which  the  middle  one 
is  the  longest ;  the  posterior  spines  are  simple ;  they  gradually  become 
smaller  and  then  disappear  entirely.  The  vulva  is  situated  behind 
the  middle  of  the  body. 

Male. — 10*5  mm.  long  by  0*6  mm.  broad.     Head  terminates  in  a 


GNATHOSTOMA    SPINIGERUM 


385 


globular  swelling  with  two  large  lips.  Neck  3  mm.  broad.  In  front  of 
neck  eight  rows  of  simple  spines  directed  backwards.  Anterior  half  of 
body  with  cuticular  laminae,  posterior  unarmed.  Two  pre-anal  and 
two  post-anal  papillae.     Bursa  wanting. 

Spicules  I* I  and  0*4  mm.  respectively. 

Leiper  considers  Gnathostonia  siainense  to  be  identical  with 
Guathostoma  spinigerum. 

The  single  specimen  described  by  Levinsen  was  found  by  Deiintzer  in 
Bangkok  (Siam),  and  was  obtained  from  a  young  Siamese  woman  who  suffered  from 
a  small  tumour  of  the  breast  which  had  developed  in  the  course  of  a  few  days. 
After  the  disappearance  of  the  tumour,  nodules  the  size  of  beans  were  found  in  the 
skin  ;  out  of  one  of  these  the  worm  was  obtained.  The  same  observer  saw  this 
affection  in  two  other  persons. 

A  closely  related  species,  Gnathostoma 
spinigerum^  Ow.,  lives  in  the  stomach  of 
wild  cat  {Felis  catus),  puma  {Felis  con- 
color),  tiger  {Felis  ti^ris),  and  domestic  cat 
(India) ;  another  species,  Gnathostoma 
hispidutn,  Fedsch.,  1839,  in  the  stomach 
of  pigs  in  Turkestan,  Annam,  Hungary, 
Congo,  and  by  Collin  in  the  stomach  of  an 
ox  (Berlin). 

Gnathostoma  sp.  in  pariah  dogs, 
Calcutta.  G?tathostoma  sp.  in  monkeys, 
French  Guiana.  They  produce  large 
fibrous    thickenings  in   the   stomach   wall. 

Gnathostoma   spinigerum, 
Owen,   1836. 

Cuticle  of  bulb  with  eight  rows 
of  chitinous  laminae  with  their 
posterior  edges  notched  into  spines. 
The  laminae  on  the  anterior  portion 
of  the  body  are  similar  trident 
laminae.       In    the    middle 'of     the 

body,  the  laminae  are; simple  and  conical,  cuticle  posteriorly  is  un- 
armed.    Mouth  with  two  fleshy  lips. 

Male  5  mm.  long  byo'S  mm. broad;  tail  spiral,four  pairs  of  papilla. 

Female  about  twice  as  long  ;  tail  straight,  trilobed. 


Fig.  275.—  Gnathostoma  siainense  :  to  the 
left,  the  entire  worm  (8/i) ;  to  the  right  the 
head  seen  from  above,  with  two  fleshy  lips 
(about  40/1).     (After  Levinsen.) 


Family.     Dracunculidae,  Leiper,  191 2. 

Genus.     Dracunculus,  Kniphoff,  1759. 

Anterior   end    rounded    with    a    cuticular   thickening    or    shield. 
Mouth  triangular  with  two  lips.     Alimentary  canal  atrophied. 
25 


THE   ANIMAL   PARASITES   OF   MAN 

Dracunculus  medinensis,  Velsch,  1674. 

Syn. :  Vena  ?nedmensis,V ehch,  1674;  Dracunculus  persarum,  Kampfer,  1694; 
Gordius  medinefists,  Linne,  1758  ;  Filaria  dracu?tculus,  Bremser,  1819  ;  Filaria 
cBthiopica,  Valenciennes,  1856;  Dracunculus  medinensis^  Cobbold,  1864;  Guinea 
worm^  Medina  worm. 

The  females  attain  a  length  of  50  to  80  cm.,  or  even  more,  and 
average  1-5  to  17  mm.  in  diameter.  They  are  whitish  or  yellowish 
in  colour.  The  anterior  extremity  is  roundish  and  bears  a  cuticular 
thickening  or  shield.  The  triangular  mouth  opening  is  surrounded 
by  two  projections  or  lips,  behind  which  on  the  shield  there  are  two 
lateral  and  four  sub-median  papillae  ;  the  posterior  end  terminates  in 
a  spine,  ventrally  directed,  and  about  i  mm.  in  length  ;  the  alimentary 
canal  below  the  oesophagus  is  atrophied,  but  not  entirely  obliterated ; 
anus  absent ;  the  lateral  lines  are  very  flat.  The  greater  part  of  the 
body  is  occupied  by  the  long  uterus,  in  which  a  great  number  of 
young  larvae  are  always  found.  The  ovaries  probably  lie  at  the  ends 
of  the  uterus  ;  the  vulva  lies  just  behind  the  cephalic  shield.  During 
parturition  the  uterus  is  prolapsed  through  this  opening. 

The  male  is  almost  unknown.  Leiper  in  an  experimentally  in- 
fected monkey  found  two  males  22  mm.  long,  one  from  the  psoas 
muscle,  the  other  from  the  connective  tissue  behind  the  oesophagus. 

Occurrence.  —  Filaria  medinensis  has  been  known  since  the  most  remote 
period.  The  "  fiery  serpents  "  that  molested  the  Israelites  by  the  Red  Sea,  and 
which  Moses  mentioned,  were  probably  filariae.  The  term  Apa/c  '^vnov  occurs  in 
Agatharchides  (140  B.C.).  Galen  called  the  disorder  dracontiasis  ;  the  Arabian 
authors  were  well  acquainted  with  the  worm.  It  is  found  not  only  in  Medina  or 
Arabia,  but  also  in  Persia,  Turkestan,  Hindustan.  The  Guinea  worm  is  also  widely 
distributed  in  Africa,  on  the  coasts  as  well  as  in  the  interior.  It  occurs  in  the  Fiji 
Islands.  It  was  carried  to  South  America  by  negro  slaves,  but  is  said  at  the  present 
lime  to  exist  in  only  quite  a  few  places  (British  Guiana,  Brazil  [Bahia])  ;  it  is  also 
observed  in  mammals  (ox,  horse,  dog,  leopard,  jackal  \Canis  lapuster\  etc.). 

Dracunculus  medinensis  in  its  adult  stage  lives  in  superficial  ulcers 
on  the  body  surface ;  it  is  seen  most  frequently  on  the  lower  extremi- 
ties, more  especially  in  the  region  of  the  ankle,  but  it  also  occurs  in 
other  parts  of  the  body — on  the  trunk,  scrotum,  perineum,  on  the 
upper  extremities,  and  in  the  eyelids  and  tongue.  Sometimes  there 
is  only  one  ulcer  and  one  worm,  but  more  commonly  several.  It 
attacks  man  without  distinction  of  race,  age  or  sex.  It  is  observed 
most  frequently  during  the  months  of  June  to  August. 

Life  history.^ — When  about  a  year  old  the  worm  seeks  the  surface 

'  The  larvae  resemble  those  of  Cticullanus  elegaus  parasitic  in  the  perch  {Perca  Jluviatilis). 
The  larv£)e  of  this  species  develop  in  Cyclops  sp.  Fedschenko  in  1870,  at  Leuckart's  suggestion, 
succeeded  in  observing  the  invasion  of  Cyclops  by  Guinea  worm  larvae.  They  penetrate  not 
per  OS  but  through  the  exoskeleton.  Newly  hatched  larvae  (in  bananas)  will  cause  infection  of 
monkeys. 


DRACUNCULUS   IIEDINENSIS 


387 


Fig.  277.  —  Anterior  ex- 
tremity of  Guinea  worm, 
showing  dorsal  and  ventral 
lips,  one  lateral  and  two  sub- 
median  papillae  and  the  lateral 
line.     (After  Leuckart.) 


Fig.  276.  —  Guinea 
worm  {Dracunculus  tnedi- 
nensis.    (After  Leuckart.) 


Fig.  278. — Dracunculus  medinensis.  a, 
anterior  extremity  seen  end  on  ;  O,  mouth  ; 
P,  papillae  ;  b,  female  reduced  more  than 
half;  er,  larvae  enlarged.     fAfler  Claus.) 


388 


THE   ANIMAL   PARASI'l  ES   OF  MAN 


of  the  body  and  produces  there  a  thickening  as  big  as  a  florin.  Over 
this  a  vesicle  forms  which  eventually  ruptures,  and  at  the  bottom 
of  the  ulcer  can  be  seen  a  hole  from  which  a  part  of  the  worm  may 
project.  On  bathing  the  sides  of  the  ulcer  with  water,  a  drop  of 
fluid,  at  first  clear  then  milky,  exudes.  This  contains  numerous  larvae. 
In  other  cases  a  thin  tube  an  inch  long  is  prolapsed  (through  the 
vulva).  This  is  probably  the  uterus,  but  the  mechanism  of  parturition 
is  not  clearly  known.  It  lasts  for  about  a  fortnight.  An  abundant 
supply  of  larvae  can  be  got  by  placing  wet  compresses  on  a  fresh 
ulcer.     In  a  few  hours  a  mass  of  larvae  is  obtained. 

The  larvae  are  500  yu<  to  750  yit  by  15/1,  to  25  yit,  with  a  long  slender 
tail  about  one-third  of  the  total  length.  The  cuticle  is  transversely 
striated.  The  body  is  flattened.  They  possess  an  oesophagus  and 
gut.     At  the  anus  there  are  apparently  glandular  structures. 

The  larvae  live 
and  move  actively  in 
water  for  about  two 
days,  the  majority 
dying  on  the  third 
(Leiper).  If  a  num- 
ber of  Cyclops  sp. 
have  been  collected 
and  isolated  in  clean 
water,  and  the  larvae 
are  now  added,  the 
further  development 
can  be  traced. 

The  larvae  enter 
the  Cyclops,  accord- 
ing to  most  authori- 
ties, by  penetrating 
the  exoskeleton,  but 
according  to  Leiper  this  is  impossible ;  they  must  enter  by  the  mouth 
and  penetrate  the  gut  in  order  to  reach  the  body  cavity.  In  eight  days 
moult  I  takes  place,  the  striated  cuticle  bemg  cast  off.  In  ten  days 
moult  2  takes  place.  In  five  weeks  the  larva  is  mature.  If  now  the 
infected  Cyclops  is  placed  in  0*2  per  cent.  HCl  solution  the  Cyclops  is 
killed  immediately,  but  the  larvae  are  stirred  into  activity,  escape  from 
the  body,  and  swim  about  in  the  acid.  This  suggests  that  infection 
in  nature  probably  takes  place  by  the  swallowing  of  infected  Cyclops ; 
Leiper,  by  feeding  Cyclops  containing  mature  larvae  to  a  monkey,  found 
in  it,  post  mortem  six  months  later,  two  immature  females  30  cm.  long 
and  two  males  22  mm.  long. 

In  certain  areas  the  new  cases  occur  principally  in  June.     Five 


Fig.  279.  —  Transverse  section  of  temale  Guinea  worm  ; 
u.,  uterus  containing  embryos  ;  i.,  intestinal  canal  ;  ^.,  ovary. 
(After  Leuckart.) 


DRACUNCULUS    MEDINENSIS 


389 


weeks  later  the  larvae  will  become  mature  in  Cyclops,  so  that  infection 
of  Cyclops  is  taking  place  in  July  or  August,  and  from  then  to  June 
about  ten  months  elapse,  giving  the  period  of  development  in  man. 

Pathology.  —  The  initial  induration  is  accompanied  by  itching. 
Urticarial  eruptions  are  described  in  Dahomey  and  Mauretanis^ 
accompanied  by  fever,  rigors,  blood-shot  conjunctiva,  and  prostration 
resembling  fungus  poisonmg.  Symptoms  last  for  one  to  two  days, 
later  the  worms  appear  on  the  surface. 

If  the  worm  is  ruptured  in  an  attempt  to  extract  it,  disastrous 
results  may  occur  through  the  escape  of  the  larvae  into  the  tissues  : 


Fig.  280. —  Cyclops  virescetis^  $.  8,  Female,  ventral  view,  x  120;  9,  anterior  antennas 
X  240;  10,  urosome  and  last  thoracic  segment,  x  240;  ii,  foot  of  first  pair,  x  320; 
12,  15,  16,  foot  of  second,  third  and  fourth  pairs,  x  240;  14,  foot  of  fifth  pair,  x  440;  13, 
last  thoracic  segment  and  first  segment  of  urosome  of  male,   x  240. 

fever,  inflammation,  abscess,  sloughing,  ankylosis,   even  death  from 
sepsis.     Eosinophilia  is  often  marked,  11  to  13  or  even  50  per  cent. 

Extraction. — (i)  The  native  method  consists  in  rolling  the  worm 
round  a  stick ;  i  in.  to  2  in.  are  extracted  each  day,  the  process 
taking  about  a  fortnight ;  (2)  Emily  used  injections  of  i  in  i,coo 
sublimate  into  the  swelling  or  into  the  worm  itself  fixed  by  a  ligature. 


390  THE   ANIMAL   PARASITES    OF   MAN 

(3)  Beclere  chloroforms  the  worm  ;  (4)  the  worm  can  be  more 
easily  removed  when  all  the  embryos  have  been  deposited  (two  to 
three  weeks). 

Cyclopidce. — Cephalothorax  ovate,  clearly  separated  from  abdomen. 
Anterior  antennae  of  female  when  bent  back  scarcely  ever  stretch 
beyond  the  cephalothorax.  The  second  antenn^e  are  unbranched. 
First  four  pairs  of  feet  two-branched,  outer  branches  three-jointed. 
The  fifth  pair  of  limbs  are  rudimentary  alike  in  both  sexes,  usually 
one-jointed.  There  is  no  heart.  The  female  has  two  egg  sacs 
containing  about  fifty  eggs. 

Genus.     Cyclops,  Miiller,  1776. 
Mandible  palp  rudimentary,  reduced    to   a   tubercle  bearing  two 
branchial  filaments.     Maxillary  palp  rudimentary  (obsolete).     Lower 
foot-jaw  non-prehensile.     Head  ankylosed  to  first  thoracic  segment. 

Family.     Filariidae. 

Sub-family.     Filarllnae. 

The  residue  after  exclusion  of  the  ArduenniiKX  and  Onchocercincv. 

Genus.    Fllaria,  O.  Fr.  Miiller,  1787. 

Very  long,  slender  Nematodes,  without  excretory  vessels  or 
excretory  pore,  the  males  of  which  are  usually  considerably  smaller 
than  the  females.  Mouth  round,  without  lips,  unarmed.  The  lateral 
lines  occupy  one-sixth  of  the  circumference  of  body.  The  tails  of  the 
males  are  bent  or  spirally  rolled,  and  bear  little  wing-like  appendages. 
The  two  spicules  are  unequal ;  almost  always  there  are  four  pre-anal 
papillae,  but  the  number  of  post-anal  papillae  varies.  The  vulva  is 
always  situated  at  the  anterior  extremity.  Parasitic  chiefly  in  the  serous 
cavities  and  in  the  subcutaneous  connective  tissue.  Insufficiently 
defined. 

Filaria  bancrofti,  Cobbold,  1877. 

Syn.  :  Trichina  cystica,  Salisbury,^  1868  {iiec  Filaria  cystica,  Rud.,  1819)  ; 
Filaria  sanguinis  hofninis,  Lewis,  1872  ;  Filaria  sanguinis  hominis  cFgyptiaca^ 
Sonsino,  1875  ;  Filaria  wiichereri,  da  Silva  Lima  ;  Filaria  sa7tgui?tis  hominum^ 
Hall,  1885;  Filaria  sanguinis  hominis  nocturna,  Manson,  1891  ;  Filaria  nocturna, 
Manson,  1891. 

These  parasites  of  man  were  for  a  long  time  only  known  in  their  larval 
stage.  They  were  discovered  in  1863  in  Paris  by  Demarquay,  in  the  hydrocele 
fluid  of  a  Havanese  emptied  by  puncture  ;  they  were  next  observed  by  Wiicherer, 
in  Bahia,  in  the  urine  of  twenty-eight  cases  of  tropical  chykiria  ;  they  were  likewise 
observed  in  North  America  by  Salisbury,  who  gave  them  the  name  of  Trichina 
cystica.  The  next  discoveries  (in  Calcutta,  Guadeloupe,  and  Port  Natal)  related  to 
rhyluria  patients,  until  Lewis  discovered  the  larvae  in  the  blood  of  man  (India),  and 

*  C.  W.  S  iles  ("American  Medicine,"  1905,  ix,  p.  682)  is  of  the  opinion  ihat  Salisbury's 
Trichina  cystica  is  idenlical  with  Oxyuris  vermicularis. 


FILARIA   BANCROFT! 


391 


found  they  were  almost  always  present  in  persons  suffering  from  chyluria,  elephan- 
tiasis, and  lymphatic  enlargements  ;  he  also,  in  exceptional  cases,  found  them  in 
apparently  healthy  persons  {Filaria  sanguinis  ho  minis).  Lewis  and  Manson  studied 
the  disease  and  the  filariae  of  the  blood  very  minutely,  and  became  aware  that  the 
filaricC  were    sucked    up   bv  mosquitoes  with   the    blood.     Manson    described   the 


Fig.  2%i.—FiIaria  bancrofti.  1,  Anterior  portion  of  male;  2,  two  rows  of  papillse  on 
head  ;  3,  papilla  of  tail  of  male  ;  4,  cloaca  of  male  showing  tips  of  spicules  and  gubernaculum  ; 
5,  the  spicules  and  gubernaculum  of  male.     (After  Leiper.) 

metamorphoses  that  take  place  within  the  body  of  the  mosquito.  The  adult 
female  was  discovered  in  Queensland  by  Bancroft,  and  soon  after  Lewis  found  it 
in  Calcutta  ;  it  was  described  by  Cobbold  as  F.  bancrofti.  The  male  was  first 
seen  by  Bourne  in  1888. 


392  THE   ANIMAL   PARASITES   OF   MAN 

Head  bougie-like,  i.e.^  separated  by  a  narrowing  from  the  neck, 
having  two  rows  of  minute  papillae.  Cuticle  has  extremely  fine 
striations. 

Female. — 50  to  65  mm.  long  by  1*5  to  2  mm.  broad.  Vulva 
0-4  to  07  mm.  behind  the  head.  Anus  about  \  mm.  from  the  tip 
of  the  tail  (vulva  i  to  1*3  mm.  from  head,  and  anusciyto  28  mm.  from 
tail  according  to  other  authors).  The  vagina'  is  a  muscular  tube  form- 
ing three  bold  loops,  and  has  terminally  a  pyriform  enlargement. 
Uterus  double  (or  single).     Ovoviviparous. 

Male. — 25  to  30  mm.  long  by  o'l  mm.  thick  (40  by  o*i  mm.  ac- 
cording to  various  authors).  Probably  two  pairs  of  pre-anal  papillae, 
eight  pairs  of  peri-anal,  two  pairs  of  post-anal  papillae,  and  one  pair 
terminal.  Tail  curved.  Two  spicules,  0*2  and  o'6  mm.  respectively, 
and  a  cup-like  gubernaculum.  The  long  spicule  is  cylindrical, 
expanded  proximally  and  tapering  distally  to  a  filament  with  wings. 
At  the  tip  it  is  spoon-like.  The  short  spicule  is  of  the  same  diameter 
throughout.  It  is  gutter-like,  coarsely  marked.  Testis  uncoiled, 
terminating  in  a  snowdrop-like  process  (Leiper). 

^^^' — 40  /^  by  25  jjb.  They  do  not  appear  to  possess  a  true  shell, 
but  only  an  embryonal  or  vitelline  membrane  secreted  by  the  ovum. 

Embryos. — In  the  posterior  part  of  the  uterus  eggs  occur,  in  the 
anterior  part  embryos;  the  larvae  at  birth  measure  127  fxio  200/1  by 
8/A  to  10  fji.  In  the  blood  they  measure  in  the  fresh  260  fju  by  7*5  fx  to  8/^. 
In  stained  films,  owing  to  shrinkage,  there  is  great  variation  in  size,  from 
154 /A  to  311 /A.  Probably  260  yu,  to  285 /a  is  the  average  in  stained 
films. 

Geographical  Distribution. — Europe  :  Two  cases  recorded,  one 
from  near  Barcelona.  The  patient  suffered  from  h^mato-chyluria 
and  enlarged  scrotum  wqth  mikrofilariae  in  the  blood.  A  second 
case  from  Siena.  Africa  :  The  filarial  index  has  not  been  estimated 
for  various  parts.     In  Nigeria  it  is  about  10  per  cent. 

//<76/7(7/.— Lymphatic  glands  :  e.g.,  inguinal,  femoral,  iliac,  lumbar, 
mesenteric,  bronchial,  superficial  cervical,  epitrochlear. 

Lymphatic  vessels  :  e.g.,  those  draining  into  the  receptaculum  chyli 
of  the  spermatic  cord,  in  the  thoracic  duct  and  in  various  different  parts. 

Organs,  etc. :  Testis,  epididymis,  spermatic  cord,  tunica  vaginalis, 
mammary  cyst,  and  in  abscesses. 

They  may  occur  in  masses,  but  usually  only  a  few  (one  to  eight). 
Females  are  commoner  than  males.  Dead  and  calcified  worms  are 
common  in  the  various  sites. 

Distribution  of  Larva:  in  Body,— ThtSQ  are  by  no  means  uniformly 
distributed,  but  occur  in  greater  number  in  tiie  capillaries  of  the 
lungs.  Besides  the  lungs  they  occur  in  the  capillaries  of  other  organs, 
as  the  following  data  of  Rodenwaldt  show:— 


Mikrofilariae 

Lungs       

...     I34,82i' 

Spleen 

Liver        

4,884 

Brain 

Kidneys 

•••       15,253 

Glands 

Glomeruli 

8,008 

Marrow 

Parenchyma 

7,245 

Blood 

FILARIA   BANCROFTI  393 

Mikrofilariae 

1,666 

3,833 

o 

o 

3>ooo 

The  following  data  of  Rodenwaldt  refer  to  the  larvae  of  Filaria 
itnmitis  m  the  dog.  They  are  commoner  in  organs  than  in  vessels, 
and  especially  in  the  capillaries  of  the  organs,  but  in  the  lungs  they 
appear  to  be  equally  distributed  in  capillaries,  arteries  and  veins. 

The  length  of  life  of  larvae  is  unknown,  but  they  appear  to  be 
destroyed  in  the  kidneys,  as  dead  calcified  specimens  are  fairly 
numerous  in  the  capillaries  of  the  vasa  recta  of  the  medullary 
substance. 

Kidneys  :  mainly  in  the  glomerular  capillaries  and  those  of  the 
vasa  recta. 

Liver  :  in  the  capillaries  of  the  portal  system,  especially  in  those 
between  the  interlobular  and  the  central  intralobular  veins. 

Periodicity  of  Larva^.^ — Roughly  speaking,  the  larvae  of  Filaria 
bancrofti  are  found  in  the  peripheral  blood  only  during  the  night, 
disappearing  (but  not  entirely)  during  the  daytime.  Their  periodicity 
and  that  of  Loa  loa  larvae  is  shown  by  the  table  on  p.  394,  based  on 
that  of  Smith  and  Rivas  (Amer.  Jouru.  Trop.  Dis.  and  Prev.  Med., 
1914,  vol.  iii,  p.  361). 

It  was  discovered  by  Mackenzie  that  this  periodicity  could  be 
reversed  by  making  the  patient  sleep  during  the  daytime,  showing 
that  the  phenomenon  was  in  some  way  dependent  on  sleep  or  its 
attendant  phenomena.  Rodenwaldt  gives  the  following  explanation 
of  the  phenomenon  of  periodicity  : — 

Mikrofilariae  come  to  rest  in  capillaries.  After  passing  up  the 
thoracic  duct  they  would  reach  the  capillaries  of  the  lungs  by  the 
superior  vena  cava.  Here  they  occur  in  immense  numbers.  In  the 
case  of  Loa  loa  larvae  (which  have  a  diurnal  periodicity)  some  of 
these  are  forced  out  by  the  increased  force  and  rapidity  of  the 
pulmonary  circulation  during  the  day,  but  are  able  to  rest  (owing 
to  their  sticky  sheath  ?)  in  the  peripheral  capillaries  on  their  way  to 
the  capillaries  of  the  organs.  During  the  night  the  force  of  the 
current  through  the  lungs  is  relaxed  and  consequently  they  are  able 

'  These  figures  refer  to  i  c.c.  of  each  organ,  and  were  estimated  by  cutting  sections  of 
definite  thickness  (30  fi  to  40  fi)  and  counting  the  filariae  in  a  definite  area  of  section,  e.^., 
I  cm.^  The  organs  before  removal  from  the  body  have  their  vessels  tied,  and  are  then  fixed  in 
hot  alcohol. 

^  For  determining  periodicity  measured  quantities  of  blood,  e.^,  20  mm.*,  should  be  used. 
A  thick  film  is  made  of  the  whole  quantity.  The  numbers  present  in  this  quantity  may  vary 
from  three  or  four  to  300  or  400. 


394 


THE   ANIMAL   PARASITES   OF   MAN 


to  remain  in  the  pulmonary  capillaries  and  do  not  appear  in  the 
capillaries  of  the  systemic  circulation.  If  it  is  true  that  the  periodicity 
of  Loa  loa  cannot  be  reversed  by  changing  the  hours  of  sleep,  then 
the  explanation  is  incomplete.  In  the  case  of  the  larv?e  of  Filar ia 
bancrofti  (which  have  a  nocturnal  periodicity),  in  order  to  apply  the 
same  explanation  we  must  further  assume  that  the  mikrofilariae  have 
less  power  of  resisting  the  force  of  the  capillary  current  {i.e.,  are  less 
sticky).  They  are  washed  out  of  the  pulmonary  capillaries  by  day 
and  by  night,  but  it  is  only  at  night,  when  the  blood-stream  in  systemic 
capillaries  is  less  rapid,  that  they  are  able  to  rest  there.  In  the  day- 
time they  are  washed  on  until  they  reach  the  capillaries  of  the  organs 
(possibly  again  the  lungs).     The  reversal  of  tlie  periodicity  by  sleeping 


Larvae  of  L. 

Average  132. 

Deviations 

fiom  average 

Case  i. 

Average  1,000 

Case  2. 

Average  1,570 

loa  in  equal 

F.  bancro/ti 

(about). 

F.  bancro/ti 

(about). 

quantities  of 

larvae  in  i  c.c. 

Deviations 

larvae  in  i  c.c. 

Deviations 

blood 

of  blood 

from  average 

of  blood 

from  average 

2  a.m.   ... 

9 

-    123 

3,500 

+  2,500 

6,500 

+  3.930 

4  a.m.   ... 
6  a.m.   ... 

1 1 

-    121 

3,200 

+   2,2O0 

5,200 

+  3.630 

41 

-      91 

2,800 

+    1,800 

2,000 

+       430 

8  a.m.   ... 

1 68 

+      36 

900 

-        100 

1,100 

-       470 

lo  a.m.   ... 

298 

+  1^6 

210 

-        790 

350 

—    1,220 

12  noon  ... 

531 

+  389 

30 

-        970 

50 

-    1,520 

2  p.m.  ... 

252 

+  120 

20 

-        980 

4C 

-    1,530 

4  pm.   ... 
6  p.m.   ... 
8  p.m.  ... 

146 

+     14 

10 

-        990 

30 

-    1,540 

91 

-     41 

40 

-        960 

40 

-    1,530 

2.1 

-     99 

60 

-        940 

ICO 

-    1,470 

lo  p.m.  ... 

5 

-  127 

600 

-        400 

800 

-        770 

1 2  midnight 

5 

-  127 

750 

-        250 

2,600 

+   1,030 

Total  ... 

1,580 

— 

12,120 

— 

18,810 

— 

during  the  daytime  admits  of  a  similar  explanation.  If  this  explana- 
tion be  true,  then  a  prolongation  of  the  day  conditions,  e.g.,  by 
continued  exercise,  should  result  in  still  keeping  the  larvae  out  of  the 
circulation,  but  this  does  not  appear  to  be  the  case. 

In  certain  countries,  e.g.,  Fiji,  Samoa,  Philippines,  West  Africa, 
larvae,  apparently  those  of  Filavia  bancrofti,  show  no  periodicity.  In 
Fiji  the  usual  intermediate  host  is  Siegoinyia  psendoscutellaris,  a  day- 
bitmg  mosquito,  so  that  possibly,  as  Bahr  suggests,  the  mikrofilari^ 
have  partly  adapted  themselves  to  the  habits  of  their  intermediate 
host,  as  the  nocturnal  mikrofilariae  are  adapted  for  transmission  by  a 
nocturnal  feeding  mosquito,  e.g.,  Culex  fatigans,  but  how  this  could 
come  about  is  a  mystery.  It  is  not  certain  in  all  cases  whether 
the  non-periodic  mikrofilariae  really  belong  to  Filaria  bancrofti; 
some  may  be  L.  loa  larvae,  or  possibly  unknown  larvae.  An  exact 
morphological  description  of  these  larvae  is  therefore  always  necessary. 


FILARIA    BANCROFTI  395 

Preservation  of  Living  Larvce. — Blood  from  the  vein  (or  finger 
puncture)  is  shaken  up  with  twenty  times  its  volume  of  sterile  0*9  per 
cent,  salt  solution,  and  kept  in  an  ice  cupboard  (F'iilleborn). 

Concentration  of  Larvce. — {a)  The  above  mixture  is  haemolysed  with 
water  and  then  sufficient  salt  solution  added  to  make  up  to  0*9  per 
cent.  The  solution  is  allowed  to  stand  or  can  be  centrifugalized. 
(6)  The  blood  is  mixed  with  sodium  citrate  and  centrifugalized  ;  the 
larvae  are  found  in  the  leucocytic  layer  (Bahr).  (c)  Allow  blood  to 
clot  in  a  small  tube  ;  the  larvae  appear  on  the  surface  of  the  clot  and 
are  so  got  in  pure  serum.  A  drop  of  blood  may  also  be  allowed  to 
clot  on  the  slide ;  the  larvae  are  found  in  the  clear  areas  of  serum. 
{d)  Haemolyse  blood  with  water  or  acetic  acid.  Centrifugalize,  make 
smears  from,  or  examine  the  sediment. 

Removal  of  Red  Corpuscles. — The  blood  film  is  allowed  to  stand  for 
some  minutes  in  a  moist  atmosphere.  The  staining  solution  is  sucked 
through  with  blotting  paper  :  the  larvae  stick  to  the  slide,  while  the 
corpuscles  are  washed  out. 

Morphology  of  Larvce. — Wet  staining:  Azur  II  one  part,  0*9  per 
cent.,  salt  solution  3,000,  or  very  dilute  Giemsa  or  ripened  methylene 
blue  or  neutral  red  solutions.  Place  a  drop  on  the  slide  and  add  a 
drop  of  blood  to  this.  The  larvae  remain  alive  for  one  or  more  days ; 
it  sometimes  takes  twenty-four  hours  to  stain  some  particular  structure. 
Differentiation  by  drawing  through  weak  eosin  solution  is  often 
useful.  This  method  is  the  best  for  finest  details.  The  excretory 
pore,  anal  pore,  excretory  cell,  and  chief  "  genital  "  cell  stain  hrst, 
then  the  matrix  cells  and  finally  the  column  of  nuclei. 

Wet  fixation  and  staining :  The  blood  is  spread  on  a  large  cover- 
glass — floated  on  the  surface  of  70  per  cent,  alcohol  heated  to  about 
70°  C.  Wash  in  water,  (i)  overstain  with  i  in  1,000  azur  II  solution, 
warming  slightly ;  (2)  differentiate  with  {a)  absolute  alcohol  (con- 
taining, if  necessary,  a  trace  of  HCl),  or  (6)  with  absolute  alcohol 
96  per  cent,  ninety  parts,  anilin  oil  ten  parts  ;  (3)  clear  in  origanum, 
bergamot  or  cajeput  oil ;  (4)  mount  in  balsam.  Or  stain  with  haema- 
toxylin,  e.g.,  Mayer's  glycerine  alumhaematein,  heating  till  slightly 
steaming.  Differentiate  with  acid  (2  per  cent.  HCl)  alcohol  if  ov^er- 
stained.     Clear  and  mount  as  above. 

Dry  fixation  and  staining:  (i)  With  azur  II  as  above,  or  (2)  with 
haematein  (warm).  Exarnine  the  dried  films  in  the  usual  way  without 
a  cover-glass.     The  azur  stains  the  excretory  and  genital  cells  clearly. 

Thick  films  :  (i)  The  blood  is  smeared  out  fairly  thickly  over  an 
area  as  big  as  a  sixpence. 

(2)  Dry  quickly  to  prevent  shrinking,  using  carefully  a  spirit  lamp 
in  a  moist  climate. 

(3)  Place  films  downwards  in  water  for  a  few  minutes. 


396  THE   ANIMAL   PARASITES   OF   MAN 

(4)  Fix  in  alcohol. 

(5)  Stain  withazur  II,  i  in  1,000.  Differentiate  as  above.  Examine 
as  a  dry  film.  This  method  suffices  for  showing  the  excretory  cell  and 
the  Gi  cell ;  or 

(6)  Stain  with  haematein  (slightly  steaming),  especially  for  the 
column  of  nuclei  and  the  sheath.  The  fixation  in  alcohol  in  this 
case  may  be  omitted. 

(7)  The  removal  of  the  haemoglobin  and  the  fixation  may  be 
combined  by  using  Ruge's  mixture  (formalin  2  per  cent.,  containing 
I  per  cent,  acetic  acid)  or  acetic  alcohol  (glacial  acetic  i,  alcohol  3).^ 

Striidnre  of  Larvcv. — (i)  Subcuticular  cells  :  By  vital  staining,  at 
intervals  underneath  the  cuticle  are  seen  a  series  of  spindle-shaped 
cells — the  sitbctiticular  matrix  cells  of  Rodenwaldt,  the  muscle  cells 
of  Fulleborn.     There  are  thirty  or  forty  or  more  of  these. 

(2)  Nerve  ring  :  Appears  as  a  break  in  the  nuclear  column  about 
20  per  cent,  of  total  length  from  the  head. 

(3)  Excretory  system  :  Consists  of  a  lateral  spherical  hollow 
excretory  pore  which  shows  a  radial  striation.  Connected  with  the 
pore  is  an  excretory  cell  which  appears  to  be  canalized.  Excretory 
pore,  29-6  per  cent,  of  length  from  head.  Excretory  cell,  30-6  per  cent, 
of  length  from  head. 

(4)  "  Genital "  cells  and  anal  pore  :  Consists  of  a  pore  opening 
ventrally  on  a  very  fine  papilla  with  which  are  connected  four 
other  cells  in  series,  the  chief  ^^  genital  "  cell  (Gi)  being  some  distance 
from  the  three  others,  which  lie  close  to  the  pore.  Gi,  70*6  per  cent., 
anal  pore,  82-4  per  cent,  of  length  from  head. 

(5)  Internal  body,  viscus,  or  reserve  material  :  Best  shown  by  vital 
staining  with  neutral  red.  This  is  agranular  strand-like  body  extend- 
ing from  527  per  cent,  to  65  per  cent,  of  length  from  head. 

(6)  Tail  end  :  (i)  Rod-like  structures  resembling  those  in  the  head, 
90  per  cent,  of  length,  (ii)  The  column  of  nuclei  extends  to  95  per  cent, 
of  length,  so  that  the  terminal  portion  is  free  from  nuclei. 

(7)  Mouth  :  Terminal  according  to  some  authors,  lateral  according 
to  others.  Some  describe  a  fang  on  the  head,  others  not.  By  vital 
staining  and  eosin  differentiation  two  rod-like  structures  with  mush- 
room-like caps  can  be  seen  behind  the  head. 

(8)  Cuticle  :  Transversely  striated.  There  is  a  longitudinal  break  in 
the  striation  on  each  side  corresponding  to  the  lateral  lines.  The  stria- 
tion is  best  shown  by  vital  staining  with  azur  II  and  eosin  differentiation. 

(9)  Column  of  nuclei  :  These  nuclei  of  the  gut  cells  form  the 
main  feature  in  ordinary  dry  films  stained  with  hematoxylin.  They 
are  separated  by  a  space  from  the  subcuticular  cells. 

'  [Acetic  alcohol  does  well  for  detecting  crescents  in  thick  films  of  malaria  blood.— 
J.  W.  W.  S.] 


FILARIA   BANCROFTI 


397 


J3  a> 


10  ^— ^ 


i 


^ 


398  THE   ANIMAL   PARASITES   OF   MAN 

Distinction  between  Mikrojilaria  bancrofti  and  Mikroloa  ha. 
Dry  Films^  HcEmatoxylin  Staining : — 

Mf.  bancrofti.  Ml.  loa. 

(i)  In  graceful  curves  (but  only  if        (i)  Kinked, 
quickly  dried). 

(2)  Tip  of  tail  free  from  nuclei.  (2)  Nuclei  extend  to  tip. 

(3)  Column  of  nuclei  separated  by        (3)  Not  so  distinctly. 

a  space  from  the  cuticle. 


(4)  Gi  cell  small,  easily  overlooked.        (4)  Gi  cell  large,  stains  deep  blue, 

cell  protoplasm  =  twice  width 
of  larva,  easily  seen. 

(5)  Excretory  cell  close  to  excretory        (5)  Excretory  cell  farther  from  pore, 

pore,  2  per  cent,  of  length.  4  per  cent,  of  length. 

Vital  Staining  with  Neutral  Red  :— 

(6)  Internal  body  or  reserve  mate-         (6)  Not  shown. 

rial  clearly  shown. 

Life  History. — In  the  stomach  of  the  mosquito  the  larvae  cast 
their  sheath  in  the  thickened  blood  in  one  to  two  hours.  In  twenty- 
four  hours  the  majority  have  reached  the  thoracic  muscles,  where 
development  proceeds.  They  are  at  first  immobile  and  of  a  ^'  sausage  " 
form  (iiOyLt  by  13ft),  with  a  short  spiky  tail.  In  three  to  five  days  the 
oesophagus  is  formed,  the  larva  now  being  0*5  mm.  long.  The  larva 
appears  to  moult  at  this  time.  After  the  gut  is  formed  papillae,  three 
or  four  in  number,  appear  at  the  tail  end.  In  two  to  three  weeks 
the  larvae  are  i'5  mm.  long.  They  now  leave  the  thorax  and  reach  the 
labium,  but  they  may  be  found  in  various  parts  of  the  body,  e.g.j  the 
legs.  They  bore  through  Button's  membrane  and  so  arrive  on  the 
surface  of  the  skin,  which  they  rapidly  enter.  Their  development  in 
man  is  unknown,  but  it  may  be  very  long,  as  children  are  not  infected 
till  4  to  5,  or  even  lOj  years  old,  but  this  may  be  due  to  unknown 
causes.  , 

Development  takes  place  in  numerous  mosquitoes.  Anophelines  : 
Myzomyia  rossii,  Pyretophorus  costalis,  Myzorhynchiis  sinensis^  Myzo- 
rhynchus  barbirostris,  Myzorhynchiis  peditceniatus. 

Culicines  :  Culex  pipiens,  Culex  fatigans,  Ciilex  skusei,  Ciilex 
gelidus,  Culex  sitiens,  Culex  albopictus,  Stegomyia  fasciata,  Stegomyia 
pseudoscutellaris,  Stegomyia  gracilis,  Stegomyia  perplexa,  Mansonioides 
imiformiSy  Mansonioidesiannulipes,  Scntomyia  alboliiieata,  Tceniorhynchtis 
domesticiis. 

Partial  development  takes  place  in  other  species. 

Pathology. — Among    the    conditions    which    Filaria    bancrofti    is 


FILARIA   BANCROFTI 


399 


400  THE   ANIMAL   PARASITES   OF   MAN 

believed  to  produce  are  lymphangitis,  varicose  glands,  especially 
inguinal  and  epitrochlear,  chyluria,  chylocele,  lymph  scrotum, 
orchitis,  abscess,  and  elephantiasis.  The  evidence  that  these 
so-called  ''filarial  diseases"  are  produced  by  F.  bancrofti  is  (i)  geo- 
graphical and  statistical  ;  (2)  pathological.  Bahr  has  contributed 
evidence  of  the  former  kind  from  his  researches  in  Fiji,  on  which 
we  may  base  the  following  statements: — 

(i)  The  prevalence  of  filarial  diseases  is  proportional  to  the 
prevalence  of  Mikrofilaria  bancrofti  in  the  blood.  Thus  in  four 
villages  examined  by  him  he  got  the  following  figures: — 


Village  A 

Village  B 

Village  C 

Village  D 

Mf.  bancrofti 

12*5  per  cent.     . 

. .     25  per  cent.     . 

31  per  cent.     . 

33  per  cent 

Filarial  diseases 

..     29        „ 

••     39 

...     58        „ 

..     34        M 

Tolal  population 

..     168 

..     114       ... 

..     425        ... 

..     222 

(2)  Out  of  257  people  with  Mf.  bancrofti  in  the  blood,  153  were 
suffering  from  filarial  diseases,  i.e.,  59  per  cent. 

(3)  Whereas  of  672  people  without  Mf.  bancrofti  in  the  blood, 
only  263  were  suffering  from  filarial  diseases,  i.e.y  37*6  per  cent. 

(4)  Again  out  of  416  people  suffering  from  filarial  disease,  153 
showed  Mf  bancrofti  in  their  blood,  i.e.,  367  per  cent. 

It  is  generally  assumed  that  all  people  suffering  from  filarial 
disease  show  at  some  (presumably  early)  stage  larvae  in  the  blood  ; 
but  we  do  not  consider  that  this  must  necessarily  be  so.  It  appears 
to  us  quite  possible  that  living  adult  filariae  may  be  present  in  the 
body,  producing  disease,  without  their  larvae  appearing  in  the  blood. 
The  absence  of  larvae  from  the  blood  in  63*3  per  cent,  of  persons 
suffering  from  filarial  disease  is,  however,  generally  explained  otherwise. 
The  adults  which  occur  in  enlarged  glands,  etc.,  get  eventually 
destroyed  by  inflammatory  reaction,  so  that  larvae  are  no  longer 
being  produced,  while  the  enlarged  gland,  etc.,  which  the  adults  have 
produced  remains.  This  explanation  assumes  that  the  larvae  of  the 
original  worm  die  in  the  circulation  or  elsewhere,  e.g.,  kidney,  but 
we  have  no  evidence  as  to  the  duration  of  life  of  larvae  in  the  human 
body;  but  also  it  assumes  that  a  person  cannot  be  reinfected  with 
filaria,  for  otherwise  there  is  no  reason  why  the  diseased  should  not  be 
infected  in  the  same  proportion  as  the  non-diseased.  But  assuming 
the  explanation  to  be  true,  it  would  explain  why  a  diseased  population 
show  larvae  in  only  about  one-third  of  the  cases.  It  must  be  borne 
in  mind  also  that  the  figures  are  rather  small. 

Pathology. — In  order  to  explain  the  effects  which  do  or  may  be 
expected  to  occur  from  obstruction  of  lymphatics,  it  is  necessary  to 
have  an  accurate  knowledge  of  the  distribution  and  connections  of 


FILARIA    BANCROFTl  4OI 

lymphatic  vessels  (and  glands)  and  the  anastomoses  of  these  vessels. 
We  can  onlv  briefly  summarize  our  knowledge  here. 

We  should  recall  also  that  considerable  destruction  or  obstruction 
of  lymphatics  or  glands  may  occui-  without  necessarily  producing  any 
lymphatic  obstruction,  at  least,  of  a  permanent  nature,  e.g.^  when  a 
mass  of  lymphatic  glands  is  destroyed  by  a  bubo  in  the  groin  or, 
again,  when  a  carcinomatous  mass  of  glands  is  removed  from  the 
axilla.  Again,  to  take  the  case  of  chyluria — where  it  is  generally 
assumed  that  obstruction  must  occur  higher  up  than  the  point  at 
which  the  intestinal  lacteals  enter  the  juxta-aortic  glands — this  disease 
may  occur,  e.g.,  in  temperate  regions,  quite  apart  from  such  obstruc- 
tion. It  is  true  that  some  of  these  cases  of  chyluria  are  not  cases  of 
chyle  in  the  urine,  but,  as  little  or  no  fat  is  present,  lymphuria. 
These  do  not  require  the  above  assumption,  but  seeing  that  true 
chyluria  may  apparently  occur  without  such  obstruction,  we  should 
be  cautious  about  explaining  this  and  other  symptoms  on  the  basis 
of  obstructions  which  theory  may  demand,  for  only  too  often  there 
are  no  post-inorteni  facts  at  our  disposal. 

Lymphangitis:  What  this  is  due  to  is  unknown.  There  is  no 
actual  evidence  of  the  occurrence  of  adults  in  the  inflamed  vessel. 
Complete  disappearance,  not  to  reappear,  of  (non-periodic)  mikro- 
filariai  from  the  blood  has  been  shown  by  Bahr  and  others  to  occur 
within  twenty-four  hours  after  an  attack  of  lymphangitis,  orchitis 
adenitis  or  simply  a  high  temperature.  This  mysterious  phenomenon 
requires  explanation.  If  the  mikrofilariae  were  being  killed  by  the 
attack,  their  dead  bodies  should  still  be  found  in  the  blood ;  or  if  the 
adults  were  being  killed,  for  all  we  know  to  the  contrary,  the  larvae 
might  well  survive.  We  consider  there  is  no  evidence  that  either 
are  affected,  but  that  for  some  reason,  as  little  understood  as  in 
periodicity,  the  larvae  now  remain  in  the  organs. 

Abscess  :  In  Fiji,  by  Bahr,  they  have  been  found  in  the  substance 
of  various  muscles,  e.g.,  quadriceps  extensor,  latissimus  dorsi,  serratus 
magnus,  in  the  popliteal  space,  groin,  axilla,  and  over  the  internal  con- 
dyle of  the  humerus,  and  in  the  upper  extremity  they  are  frequently 
infected  with  cocci.  They  not  infrequently  contain  fragments  of 
dead  adult  filarial.  Their  mode  of  origin  is  not  clear.  They  form 
nearly  30  per  cent,  of  cases  of  filariasis  in  Fiji.  Of  95  cases,  41 
showed  mikrofilariae    in  blood,  54  did  not. 

Hydrocele  and  enlarged  testis  :  In  Fiji  they  form  about  10  per 
cent.  (36  out  of  343)  of  cases  of  filariasis.  The  fluid  is  usually  sterile  ; 
mikrofilariae  were  present  in  the  fluid  in  i  out  of  it  cases.  In  the  wall 
numerous  calcified  adult  filariae  may  be  found.  The  walls  consist 
chiefly  of  hypertrophied  muscle  with  fibrous  tissue,  dilated  blood- 
vessels and  lymphatics,  the  lining  epithelium  of  which  appears  to  be 
26 


402  THE    ANIMAL    PARASITES    OF    MAN 

absent  ;  of  38  cases  14  had  mikrofilariae  in  the  blood,  24  had  not. 
Most  of  the  cases  are  associated  with  elephantiasis  of  the  scrotum 
(11  out  of  12  cases). 

Enlarged  glands  form  over  40  per  cent.  (153  out  of  343)  of  cases 
of  filariasis,  so  that  they  are  the  commonest  expression  of  filariasis 
met  with  in  Fiji.  The  glands  are  enlarged,  fibrotic,  and  the  trabeculae 
are  thickened.  The  lymphatics  are  thickened  or  represented  merely 
by  fibrous  tissue.  The  gland  also  shows  dilated  blood-vessels  and 
numerous  spaces  filled  with  lymph.  Giant-cells  are  common  in 
those  glands  which  contain  remnants  of  filariae.  Masses  of  lympho- 
cytes enclosed  by  inflammatory  or  fibrous  tissue  are  common. 
Eosinophile  cells  are  also  extremely  common,  not  only  in  the  fibrous 
tissue  of  the  glands,  but  in  other  inflammatory  or  fibrotic  conditions  : 
in  other  organs  living  or  calcified  filariae  are  "  usually "  present. 
Only  about  33  per  cent,  show  mikrofilariae  in  the  blood.  The 
epitrochlear  gland  is  frequently  enlarged  in  Fiji. 

Breinl  has  examined  enlarged  glands  and  finds  loose  vascular 
fibrous  tissue  with  lymphocytic  invasion.  In  parts,  the  lymphocytes 
collect  into  areas  200  //,  to  800  /i  in  diameter.  The  lymph  tissue 
surrounding  the  spermatic  cord  showed  abundance  of  vessels  — 
<i>  large,  (2)  small.  The  large  had  thick  walls  and  wide  lumina.  In 
other  cases  the  lumina  were  nearly  filled  by  a  thrombus  of  newiy 
formed,  fine,  loose  connective  tissue. 

Varicose  glands:  In  about  7  per  cent.  (24  out  of  343  cases)  of 
filariasis,  mikrofilariae  are  found  in  the  blood  in  50  per  cent.  (12  out 
of  24). 

Elephantiasis.— E\eph3.ni'i2isih  scroti  is  associated  with  hydrocele 
in  50  per  cent,  of  cases  (12  out  of  23) ;  in  65  per  cent,  of  cases  (15  out 
of  23)  there  are  associated  enlarged  glands  in  one  or  both  groins, 
though  also  hydrocele  and  enlarged  glands  occur  without  elephantiasis 
scroti.  In  13  out  of  27,  i.e.,  about  50  per  cent.,  cases  of  elephantiasis 
in  various  regions,  no  associated  enlargement  of  glands  is  found. 
Elephantiasis  forms  in  Fiji  less  than  10  per  cent,  of  cases  of  filariasis. 
Mikrofilariae  are  present  in  the  blood  in  36  per  cent.  (12  out  of  33) 
of  cases. 

C/^y/z/r/a.— Exceedingly  rare  in  Fiji.  Theory  would  demand  an 
obstruction  above  the  point  t)f  entry  of  the  lacteals,  viz.,  the  pre-aortic 
lymphatic  glands,  but  in  cases  in  temperate  regions  it  may  occur 
without  any  such  lesion.  In  some  of  these  cases  the  fluid  is  not 
chyle  (fat  absent),  but  presumably  lymph.  A  discussion  of  the  mode 
of  production  of  chyluria,  lymph  scrotum,  elephantiasis,  etc.,  is  at 
present  premature  ;  theory  has  far  outrun  fact.  Too  much  stress  had 
been  laid  on  the  mechanical  action  of  the  worms  to  the  almost  total 
exclusion  of  their  (or  possibly  their  larval)  toxic  action.     The  above 


FILARIA   DEMARQUAYI  4O3 

analysis  has  been  made  in  the  hope  of  acquiring  more  extended 
observations  similar  to  those  made  by  Bahr. 

Geographical  Distribution. — Filaria  hancrofti  is  known  in  nearly  all 
tropical  countries.  It  occurs  in  India,  China,  Indo-China,  Japan, 
Australia,  Queensland,  the  Islands  of  Polynesia  (with  the  exception 
of  the  Sandwich  Islands),  Egypt,  Algeria,  Tunis,  Madagascar,  Zanzi- 
bar, Sudan,  etc.,  the  south  of  the  United  States  of  America,  Brazil, 
the  Antilles,  etc.  Whether  it  is  the  same  species  in  all  cases  is 
questionable. 

Filaria  demarquayi,  Manson,  1895. 
Syn.  :  F.  ozzardi,  Manson,  1897. 

The  adult  female  F.  demarquayi  measures  from  65  to  80  mm. 
in  length  by  0*21  to  0*25  mm.  in  breadth.  The  head  has  a  diameter 
of  from  0*09  to  0*1  mm.  The  mouth  is  terminal.  The  genital 
pore  opens  at  076  mm.  from  the  head.  The  alimentary  canal  is 
nearly  straight  and  terminates  in  an  anus,  which 
is  subterminal.  The  opening  of  the  anus  is 
marked  by  a  slight  papilla.  The  tail  is  curved. 
It  rapidly  diminishes  in  size  just  below  the  anal 
papilla.  A  characteristic  pair  of  fleshy  papillae 
project  from  the  tip  of  the  tail.  The  diameter  near 
the  tip  of  the  tail  before  its  termination  is  0*03  mm. 
F.  deniarqtiayi  is  a  thicker  worm  than  Ac.  perstans. 
It  differs  from  F.  hancrofti  in  the  greater  size  of 
the  head,  in  the  smaller  tail,  and  particularly  in  the       F1G.284.— /^.^<fwar- 

,         .        ^    ,  .,    -^  _,  quayi  :    tail,    showing 

marked  fleshy  papillae  at  the  tip  of  the  tail.  These  paired  large  fleshy 
papillae  are  knobby,  and  not  simply  cuticular  as  P^P^"*-  (After Leiper.) 
in  Ac.  perstans. 

The  male  of  Filaria  deniarqnayi  has  still  to  be  found. 

The  adult  female  form  of  F.  demarquayi  was  found  by  Dr.  Galgey 
in  the  body  of  a  native  of  St.  Lucia  in  whose  blood  the  larvae  had 
been  found  during  life.  Five  adult  females  were  found  in  the  con- 
nective tissue  of  the  mesentery. 

The  larva  measures  200  //,  in  length  by  5  //,  in  breadth  ;  it  is  sharp- 
tailed,  and  has  no  sheath.  Its  movements  are  very  active,  and  the 
absence  of  a  sheath  enables  it  to  glide  along  freely  all  over  the  slide. 
It  observes  no  periodicity,  being  present  in  the  peripheral  circulation 
both  by  day  and  by  night.  As  a  rule,  some  eight  or  ten  parasites  are 
found  in  an  ordinary  preparation.  Sometimes  hundreds  of  these 
larval  hlariae  may  be  counted  on  every  slide. 

The  intermediate  host  has  not  been  discovered. 

Geographical  Distribution. — St.  Vincent,  Dominica,  Trinidad,  and 
St.  Lucia  (West  Indies),  British  Guiana,  New  Guinea  (?). 
26- 


404 


THE   ANIMAL   PARASITES   OF   MAN 


Mf.demarqua^i 


Fig.  285. — Mf.  demarquayi  in  thick  film,  dried  and  stained  with  hsematoxylin. 
6,  urshrtinken  ;  7,  shrunken.      X  1,000.     (After  Fiilleborn.) 


Filaria  taniguchi,  Penel,  1905. 

Female  68  by  0*2  mm.  in  breadth.  Cuticle  non-striated.  Mouth 
two  pairs  of  papillae.  Anus  23  mm.  from  extremity.  Vulva  1*3  mm. 
from  mouth.  Larva  164  /i,  by  8  //,,  sheathed.  Tail  truncated. 
Periodicity  nocturnal. 

Habitat. — Lymphatic  glands  of  man.     Japan. 

Filaria  (?)  conjunctivae,  Addario,  1885. 

Syn.  :  Filaria  peritonei  hontinis^  Babes,  1880;  Filaria  inermis^  Grassi,  1887  ; 
Filaria  apapillocephata^  Condorelli-Francaviglia,  1892. 

The  female  only  of  this  species  is  known.  It  measures  16  to 
20  cm.  in  length  and  0*5  mm.  in  breadth,  and  is  of  a  whitish  or 
brownish  tint.  The  cuticle  is  striated  with  fine  transverse  and  more 
marked  longitudinal  striae  with  the  exception  of  a  small  field  sur- 
rounding the  mouth,  which  is  terminal  and  has  neither  papillae  nor 
lips.  The  oesophagus  measures  o'6  mm.  in  length.  The  anus  is  3  mm. 
in  front  of  the  rounded  posterior  extremity,  and  behind  it  there  are 
two  (glandular  ?)  sacs.  The  vulva  is  close  behind  the  oral  aperture  ; 
the  vagina  soon  divides  into  two  convoluted  uteri,  which  are  filled 
with  eggs  and  embryos.     Embryos  350  yu,  by  5*5  //,. 


FILARIA  (?)   CONJUNCTIV.'E 


^05 


Fig.  286. — Filaria{})  conjiinctivce  :  to  the  left,  life  size;  to  the  light,  the 
anterior  extremity  magnified.     (After  Addario.) 

This  species  (115  mm.  long)  was  first  observed  in  Milan  by  Dubini  in  the  eye  of 
a  man;  subsequently  it  was  observed,  encysted  and  calcified  (190  mm.  long),  by 
Babes  in  the  gastro-splenic  omentum  of  a  woman  in  Budapest,  and  finally  one 
(95  mm.  long)  was  extracted  by  Vadela  from  a  tumour  the  size  of  a  pea  in  the  ocular 
conjunctiva  of  a  woman  in  Catania  (Sicily),  which  case  has  been  described  by 
Addario.  Possibly  Agamofilaria  palpebralis^  Pace,  1867  {nee  Wilson,  1844),  and 
A.  oculi  humani,  v.  Nordm.,  1832,  are  the  same  species. 


Fig.  287. — Filaria  (?)  conjunctiva  :  anterior  end  greatly  magnified  ; 
the  mouih  with  the  pharynx  in  the  middle  ;  in  the  cuticle  on  the  right 
side  the  opening  of  the  vagina,  and  behind  it  the  excretory  pore. 
(After  Grassi.) 


406  THE   ANIMAL   PARASITES   OF   MAN 

Filaria{?)  conjunctivce  is  certainly  only  an  incidental  parasite  of 
man ;  the  horse  and  ass  are  its  normal  hosts,  but  it  is  not  common 
in  these  animals,  or  is  frequently  confused  with  Haimilaria  equi, 
Gmelin,  1789. 

Group.     Agamofilaria,   Stiles,   1906. 

Not  a  generic  but  a  group  name  for  immature  Filariida'  the  deve- 
lopment of  which  does  not  admit  of  generic  determination. 

Agamofilaria  georgiana. 

Adult  unknown,  length  from  32  to  53  mm.  Maximum  diameter 
560  fM  to  640  fjL.  Head  no  cephalic  cone.  Mouth  small,  circular, 
surrounded  by  six  papillae  (two  small  latero-median  and  four 
sub-median).  The  larger  papillae  are  24  /x  from  base  to  tip.  Excre- 
tory pore  about  0*5  mm.  from  head.  Anus  64  //,  to  128  //,  from 
tip.  Cuticle  fine  striae  near  anus,  occasionally  elsewhere.  Lateral 
lines  clearly  marked.  QEsophagus  2-5  to  2-9  mm.  Rectum  200  fi 
long. 

Habitat. — Superficial  sores  on  the  ankle  of  a  negress,  Georgia, 
U.S.A. 

Agamofilaria  palpebralis,  Pace,  1867  {nee  Wilson,  1844). 
100  by  1*5  mm.,  removed  from  a  cyst  in  the  left  upper  eyelid  of 
a  boy  by  Pace,  in  Palermo. 

Agannofilaria  oculi  humani,  v.  Nordmann,  1832. 

Syn.  :  Fiiaria  lentis,  Diesing,   185 1. 

The  sexless  Nematodes  observed  in  the  lens  of  the  human  eye 
were  termed  Fiiaria  oculi  hmnani.  Only  three  cases  are  known. 
V.  Nordmann  observed  very  small  round  worms  in  the  lens  of  a 
man  and  woman  with  cataract,  and  Gescheidt  once  found  three 
specimens  in  the  lens  of  a  woman  similarly  affected. 

The  demonstration  of  nematode-like  formations  in  the  vitreous 
remains  uncertain  even  when  movements  are  observed,  and  when 
they  cannot  be  extracted  and  examined  microscopically  the  doubt 
may  occur  that  one  may  have  mistaken  the  remains  of  the  hyaloid 
artery  for  a  worm,  w^hich  it  resembles  in  form,  size  and  colour;  the 
slightest  movement  of  the  eye  also  causes  it  to  move  so  that  it 
simulates  a  living  organism. 

Accordingly  it  would  be  more  correct  to  exclude  all  the  cases  known  only 
ophthalmoscopically  (Quadri,  1857  ;  Fano,  1868  ;  Schoeler,  1875  ;  Eversbusch,  1891). 
There  then  remains  only  one  positive  case,  described  by  Kiihnt  in  1891.  In  this 
case  it  was  possible  to  follow  the  gradual  growth  of  the  parasite  for  some  time,  and 
the  worm,  which  measured  only  o"38  mm.  in  length,  was  finally  extracted. 


SETARIA  407 

Agamofilaria  labialis,  Pane,  1864. 

The  parasite  measures  30  mm.  in  length  ;  the  anterior  extremity 
is  pointed  ;  the  terminal  oral  aperture  is  surrounded  by  four  papillae ; 
the  anus  opens  0*5  mm.  in  front  of  the  posterior  extremity  ;  the  vulva 
is  2 '5  mm.  in  front  of  the  anus ;  the  uterus  is  double  ;  the  anterior  one 
passes  with  convolutions  forward  to  the  cephalic  end ;  the  posterior 
one  is  directed  backwards  and  remains  rudimentary. 

Extracted  from  a  small  pustule  on  the  inner  surface  of  the  upper  lip.  Also 
found  in  Naples  by  Pierantoni  in  1908. 

The  position  of  many  of  these  worms  is  doubtful,  and  still  more  so 
is  that  of  many  other  imperfectly  described  "  Filariae,"  which  are 
hardly  more  than  useless  and  confusing  names.  These  include  the 
following  : — 

Filaria  (?)  romanorum-orientalis,  Sarcani,  1888. 

Observed  in  the  blood  of  a  Roumanian  woman  ;  i  mm.  in  length, 
0*03  mm.  in  breadth  ;  tail  end  pointed,  a  tongue-like  appendage  on 
the  head.  Eggs  the  size  of  a  red  cell  with  developed  embryo, 
apparently  viviparous. 

Filaria  (?)  kilimarae,  Kolb,  1898. 

Several  female  specimens,  10  to  20  cm.  long  by  0*5  to  i  mm.  broad, 
were  once  found  free  in  the  abdomen  of  a  fallen  Kitii  warrior; 
according  to  Spengel,  who  examined  them,  the  oral  papillae  of  these 
worms  were  similar  to  those  of  Dracunculus  medineiisis.  Moreover, 
Kolb  classifies  together  Nematodes  that  probably  have  no  connection 
with  each  other. 

Filaria  (?)  sp.  ? 

Cholodkowsky  calls  attention  to  Filariae  that  are  still  unknown 
w^hich  cause  tumours  resembling  whitlows  on  the  fingers  of  peasants 
of  the  Twer  Government. 

Mikrofilaria  powelliy  Penel,  1905.     In  Bombay.  • 

Mikrofilaria  philippinensiSy  Ashburn  and  Craig,  1906.  In  the 
Philippines. 

Genus.     Setaria,  Viborg,  1795. 
Syn.  :  Hamularia,  Treutler,  1793  ;  Tentacidaria^  Zeder,  1800  {nee  Bosc,  1797). 

Mouth  with  projecting  peribuccal  armature  deeply  notched  on  the 
lateral  margins,  less  so  dorsally  and  ventrally.  Tail  in  both  sexes 
with  peculiar  caudal  appendages. 

Parasitic  in  serous  cavities,  especially  of  ruminants. 


4o8 


THE   ANIMAL   PARASITES   OP^   MAN 


Setaria  equina,  Abildg.,   1789. 

Syn.  :  Gordius  equinus,  AhWd^.,  1789;  Filaria  egtii,  Gm^Xwi,  1789;  Hamularia 
lymphatica^  Treutler,  1793  5  Tentacularia  subcompressa,  Zedder,  1800  ;  Filaria 
papulosa^  Rud.,  1802;  Filaria  hominis  bronchialis,  Rud.,  18 19;  Filaria  hominis^ 
Dies.,  185  r  ;  Strongylus  bronchialis,  Cobb.,  1879. 

The  body  is  whitish,   fiHform,  pointed  posteriorly.     The  cuticle 

-presents  a  delicate  transverse  striation.     The  mouth  is  small,  round, 

and  surrounded  by  a  chitinous  ring,  the  border  of  which  carries,  at  the 

sides,  two  semilunar  lips,  and  there  is  on  the  dorsal  as  well  as  on  the 

ventral  surface  a  papilliform  process ;  on  the 
tail,  corresponding  with  each  sub-median 
line,  is  a  conical  papilla.  The  male  mea- 
sures 6  to  8  cm.  in  length  ;  the  posterior 
extremity  ends  in  a  corkscrew  spiral ;  there 
are  on  each  side  four  pairs  of  pre-anal  and 
four  or  five  post-anal  papillae  ;  the  spicules 
are  unequal.  The  female  measures  9  to 
12  cm.  in  length  and  is  viviparous ;  the 
embryos  measure  0*28  mm.  in  length  and 
0*007  i^J^-  ii^  breadth. 


imsm 


Fig.  288.  —Setaria 
equina  :  left,  male ; 
right,  female.  Natural 
size.     (After    Railliet. 


Fig.  289. — Setaria  equina :  anterior  end,  magnified. 
(After  Railliet.) 


Setaria  equina  is  a  frequent  parasite  of  horses  and  asses  ;  it  inhabits  the 
peritoneal  cavity,  and  from  there  occasionally  invades  the  female  genitalia  or  even 
the  liver;  it  is  found  more  rarely  in  the  pleural  cavity  or  in  the  cranium.  The 
statement  that  it  also  occurs  in  the  subcutaneous  connective  tissue  is  probably  due 
to  confusion  with  Setaria  {Filaria)  hcemorrhagica,  Raill.,  1885  {Filaria  multi- 
papillosa,  Cond.  et  Drouilly,  1878).  Setaria  labiata  papulosa  (immature  form) 
occurs  in  the  eye  of  the  horse,  adults  in  the  peritoneal  cavity. 

Treutler,  in  1790,  found  a  filaria  in  the  enlarged  bronchial  lymphatic 
gland  of  a  patient  suffering  from  phthisis.     It  measured  26  mm.  in 


LOA  409 

length  and  had  two  spicules,  which  Treutler  mistook  for  mouth  hooks, 
hence  the  name  Hamularia.  Blanchard  mentions  another  case  from 
Geneva,  Brera  a  third  and  v.  Linstow  a  fourth.  As  shown  by  the 
synonyms,  a  few  authors  consider  this  form  to  be  a  distinct  species, 
which  is  hardly  probable. 


Genus.     Loa,  Stiles,  1905. 
Characterized  by  the  possession  of  cuticular  bosses  in  both  sexes 
(fig.  294). 

Loa  loa,  Guyot,    1778. 

Syn. :  Filaria  oculz,  Gerv.  et  v.  Ben.,  1859;  Dracunculus  oculi^  Diesing,  i860  ; 
Dracunculus  ha,  Cobbold,  1864;  Filaria  subconjunctivalis,  Guy  on,  1864. 

The  male  measures  25  to  35  mm.  in  length,  and  0*3  to  0*4  mm.  in 
breadth  ;  the  cuticle  is  not  striated,  but,  with  the  exception  of  the 
anterior  and  posterior  extremities  (r5  mm.),  is  beset  with  numerous 
irregularly  distributed  bosses  (4  /x  to  12  //,  high  by  12  yu,  to  27  //, 
broad).  The  anterior  extremity  is  somewhat  attenuated,  and  in  front 
is  conical  and  transversely  truncated.  At  the  anterior  limit  of  the 
conical  part  is  a  small  papilla  corresponding  with  the  dorsal  and  ventral 
median  lines,  and  a  little  in  front  six  non-projecting  sensory  papillae 
(two  lateral,  four  sub-median).  Excretory  pore  0*65  mm.  from  the 
anterior  end.  The  posterior  extremity  is  attenuated  and  somewhat 
curved  ventrally ;  the  anus  is  0*082  mm.  distant  from  the  rounded 
posterior  border.  In  front  of  the  anus  on  each  side  are  three  globular 
and  pedunculated  papillae  of  different  sizes,  set  close  one  behind  the 
other  but  asymmetrically  ;  behind  the  anus  on  either  side  are  two 
smaller  papillae  of  a  different  shape  ;  the  anterior  one  resembles 
the  pre-anal  papillae  in  form,  but  is  smaller ;  the  posterior  one  is 
conical,  and  rests  with  a  broad  base  on  the  cuticle.  The  spicules 
are  o'ii3  and  0'i76  mm.  long. 

The  female  measures  45  to  63  mm.  in  length  by  0*5  mm.  in 
breadth.  It  is  also  beset  with  irregularly  distributed  bosses,  which  in 
places  lie  close  to  each  other,  and  extend  to  the  anterior  extremity ; 
posteriorly  they  become  less  frequent,  but  are  not  entirely  absent. 
The  anterior  extremity  is  conical,  the  posterior  one  straight,  attenuated, 
rounded  off,  0-17  mm.  from  the  anus.  The  uteri  contain  eggs  in  the 
most  various  stages  of  development,  as  well  as  hatched-out  larvae, 
253  /x  to  262  yLt  in  length  and  47  /x  to  5  //,  in  breadth.  The  vulva  lies 
about  2  mm.  from  the  head  end.  The  vagina,  9  mm.  long,  divides 
into  two  branches,  which  at  first  ru:i  posteriorly  and  parallel  to  one 


4IO 


THE   ANIMAL   PARASITES   OF   MAN 


another  for  about  i8  mm.  One  then  bends  forward,  runs  as  far  as 
the  oesophagus,  bends  here  again  and  runs  backward  to  end  at  the 
point  of  its  first  bendmg.  The  other  branch  at  first  runs  straight 
backward  and  then  bends  forward,  but  before  reaching  the  point  of 
the  first  bend  of  the  anterior  tube  bends  backward  again,  forms 
again  a  loop  and  ends  at  the  level  of  the  anus.     The  tubes  consist  in 


Fig.  290.  —  Loa  loa :  the 
anterior  end  of  the  male,  mag- 
nified.    (After  R.  Blanchard.) 


Fig.  291. — Loa  loa:  anterior 
portion  of  the  female  as  far  as 
vulva.     (After  Loos«.) 


Fig.  2^2.— Loa  loa  in  situ. 
Natural  size.  (After  Fiille- 
born  and  Rodenwaldt.) 


Fig.  293.— Z^a  loa  :  male 
on  the  left,  female  on  the 
right.      X2.     (After  Looss.) 


the  main  of  the  uterus,  then  a  club-shaped  swelling,  the  receptaculum 
seminis,  then  the  oviduct  2  mm.  long,  and  finally  the  ovary. 

Unsegmented  eggs  measure  32  fi  by  17  fj,,  in  the  morula  stage 
40  /x  by  25  fi,  and  when  containing  embryos  50  fi  by  25  fi.  The 
vitelline  '^  shell"  of  the  egg  is,  according  to  most  authors,  stretched  by 
the  embryo  and    becomes  the  sheath   of  the    hatched  larva.     While 


LOA    LOA 


Fig.  294. — Loa  loa :  on  the  left,  the  hind  end  of  a  male ;  on   the  right,  of   a  female. 
Note  the  cuticular  bosses  shown  in  the  figure  of  the  female,      x  285.     (After  Looss.) 


(^ 


Fig.  295. — Loa  loa  :  lateral  view  of  tail  of  male  showing  Fig.  296. — Loa  loa.    a,  ventro-lateral 

papillae.     (After  Lane  and  Leiper.)  aspect  of  tail  showing  papillae  and  one 

spicule ;  b  and   c,   terminations   of  the 
two  spicules.     (After  Leiper.) 


412  THE   ANIMAL   PARASITES    OF   MAN 

still  in  the  vulva,  the  larva  measures  217  /a  to  274  /x  (average  246  /x)  in 
fresh,  146  to  226  /x  (average  192  fi)  stained. 

Site  of  Worms. — In  various  localities;  under  the  muscular  aponeu- 
roses on  extensor  surfaces  of  arms  and  legs,  lingers,  trunk,  eyelid, 
conjunctiva,  fraenum  linguae,  penis,  pericardium,  anterior  chamber  of 
eye,  and,  according  to  some  authorities,  in  lymphatic  vessels,  e.g., 
those  of  spermatic  cord.  As  many  as  thirty  adults  may  be  found. 
The  worms  appear  to  be  frequently  immature,  and  it  has  been  stated 
that  worms  in  superficial  parts  are  immature,  those  situated  deeply 
are  mature,  but  the  data  are  few. 

The  first  accounts  of  Loa  loa  —  long  since  forgotten  —  were  reported  by 
Pigafetta,  and  are  contained  in  a  book  of  travels  on  the  Congo  printed  in  1598. 
In  an  accompanying  illustration  is  depicted,  not  only  the  ancient  method  of 
extraction  of  the  Medina  worm,  but  also  the  operative  removal  of  the  filaria  from 
the  conjunctiva.  Subsequently  the  presence  of  the  worm  in  negroes  was  confirmed 
by  Bajon  in  Guiana  (1768)  and  by  Mongin  in  Mariborou  (San  Domingo),  likewise  in 
a  negro  (1770).  At  about  this  time  a  French  ship's  doctor,  Guyot,  was  cruising  on 
the  West  Coast  of  Africa  ;  he  observed  the  parasite  termed  "loa"  by  the  natives, 
and  learned  that  it  was  frequent  in  the  negroes  of  the  Congo  district.  Since  that 
time  numerous  observations  have  been  reported.  It  was  formerly  common  in  South 
America,  where  the  parasite  was  imported  by  slaves,  but  it  disappeared  when  the 
traffic  ceased  ;  it  was  particularly  prevalent  in  the  Congo,  where  it  occurs  not  only 
in  natives,  but  also  in  Europeans.  During  recent  times  it  has  repeatedly  been 
observed  in  Europe  in  negroes  as  well  as  in  white  men  who  have  lived  on  the  West 
Coast  of  Africa. 

Nematodes  of  different  size  have  been  repeatedly  observed  in  the 
eye  of  man,  in  the  anterior  chamber,  lens  and  vitreous.  For  example, 
Mercier,  in  177 1  and  1774,  removed  a  (ilaria  out  of  the  anterior 
chamber  of  two  negroes  in  St.  Domingo.  One  was  36  mm.  long. 
Barkan,  in  1876,  in  San  Francisco,  removed  one  from  the  eye  of  an 
Australian.  Again,  Cappez  and  Lacompte,  in  Brussels,  in  1894, 
observed  for  some  weeks  immature  Nematodes  in  the  eye  of  a  negro 
girl,  aged  2\  years,  and  then  removed  them.  What  these  Nematodes 
actually  were  in  these  cases  it  is  impossible  to  say. 

Structure  of  Larvct. — In  dried  films  the  larva  varies  in  size  from 
140-5  iL  to  i66*5  /^,  average,  152-5 /a;  while  another  set  of  measure- 
ments gave  the  values  isiyitto  150 /-t,  average,  143-6.  In  films  fixed 
with  hot  alcohol  the  dimensions  were  208  yLt  to  254  //,,  average,  231  yu,. 

The  nerve  ring  21-4  to  218  per  cent.  Excretory  pore  30-4  to  31-8 
per  cent.  Excretory  eel  34-8  to  37-3  per  cent.  Gi  cell  68-2  to  68*5 
per  cent.  Anal  pore  81*6  to  82-4  per  cent  of  total  length.  For  other 
details  cf.  Filaria  hancrofti. 

Larvce  in  Blood, — These  from  their  diurnal  periodicity  are  known 
as  Mikrofilaria  diuriia.  The  evidence  that  these  larvce  are  the  young 
of  the  adult  worm  Loa  loa  is  :  (i)  They  are  identical  in  structure  with 


LOA    LOA 


413 


larvae  taken  from  the  uterus  of  L.  loa  ;  (2)  their  geographical  distri- 
bution is  the  same  as  that  of  L.  loa  ;  (3)  they  eventually  occur  in 
the  blood  of  patients  suffering  from  Calabar  swellings,  a  condition 
due  to  L.  loa.  Their  occurrence  in  the  blood  in  this  latter  condition 
and  in  L.  loa  infections  we  shall  consider  later. 

Periodicity. — Here,  as  in  the  case  of  the  larvae  of  Filaria  bancrofti, 
the  larvae  that  appear  in  the  blood 
are  probably  the  overflow  simply  of 
the  larvae  which  we  assume,  on 
analogy,  to  have  their  principal  site 
in  the  lungs.  They  appear  in  the 
blood  about  the  time  of  getting 
up,  6  to  8  a.m.  (10  in  20  mm.^), 
at  12  noon  there  are  twenty-four, 
at  8  p.m.  the  number  has  fallen  to 
eighteen,  and  at  midnight  to  one, 
while  from  2  a.m.  to  6  a.m.  none, 
or  one  only,  may  be  found.  This 
periodicity  is,  as  a  rule,  a  very 
constant  one,  but  there  are  excep- 
tions, and  in  certain  cases  more 
have  been  found  at  midnight  than 
at  9  a.m.  The  periodicity  is  also 
lost  in  pathological  conditions, 
e.g.,  sleeping  sickness  {vide  also 
under  Filaria  hancrofti).  The 
possibility  of  non-periodic  Loa  loa 
larvae  should  also  be  considered. 

Pathology. — The  parasite  wanders 
about  the  body,  and  may  be  seen 
under  the  skin  in  thin  parts.  Their 
advance  is  in  some  cases  at  the  rate 
of  an  inch  in  two  minutes.  During 
their  progress  they  give  rise  to 
creeping  sensations  and  to  a  con- 
dition of  transient  oedematous  areas 
known  as  Calabar  swellings  on 
various  parts  of  the  body,  e.g.y  arm. 
These  vary  in  diameter  from  i   to 

10  cm.,  and  often  shift  their  position  an  inch  or  so  a  day.  They  give 
rise  to  a  certain  amount  of  redness,  tension  and  heat,  and  their  develop- 
ment is  promoted  by  muscular  action  of  the  part.  They  disappear  to 
reappear  elsewhere.  The  condition  is  associated  with  a  high  eosino- 
philia,  50  per  cent,  being  not  uncommon.     Patients  known  to  harbour 


Fig.  297. — Mf.  loa :  in  thick  film,  dried 
and  stained  with  hsematoxylin.  x  i,ooo. 
(After  Flilleborn.) 


414 


THE   ANIMAL   PARASITES   OF   MAN 


L.  loa,  e.g.,  native  children,  frequently  show  no  larvae  in  their  blood, 
but  they  may  do  so  after  years  of  infection.  Again,  in  patients  having 
an  infection  of  Mikrofilaria  diurnal  there  is  frequently  at  the  time  no 
evidence  of  the  presence  of  Loa  loa  adults.  Here  again  they  may 
appear  later,  but  the  conditions  which  determine  whether  persons 
infected  with  L.  loa  show  larvae  in  the  blood,  or  persons  infected 
with    Mikrofilaria   diurna    also   show    L.   loa,    are   unknown,   though 

explanations  unsupported  by  facts 
abound.  Likewise  also  the  mode 
of  production  of  the  swellings  is 
unknown. 

Not  uncommonly  Mikrofilaria 
Persians  occurs  in  the  blood  to- 
gether with  M.  diiirna. 

Duration  of  Life. — This  is  long, 
as  some  cases  have  been  observed 
five  to  six  years  after  leaving 
Africa.  The  incubation  period 
is  about  a  year. 

Life-history. — Development  of 
the  larvae  takes  place  in  the  salivary 
gl  inds  of  Chrysops  sp.  as  shown 
by  Leiper. 

Geographical  Distribution.  — 
West  Africa,  especially  in  Congo. 

Genus.     Acanthocheilonema, 

Cobbold,  1870. 

Cuticle  striated  lougitudin- 
ally.  (Esophagus  divided  into 
two  portions.  Tail  in  both  sexes 
with  short  lateral  conical  cuticular 
appendages.  Spicules  unequal, 
Fig.  2^%.  —  Acanthocheilonema  Persians,  the  larger  membranous  distally, 
I,  tail  of  male;  2,  tail  showing  cuticular    the    smaller    hooked.     Vulva    in 

flaps     devoid     of     fleshy    contents.       (After 

Leiper.)  Oesophageal  region. 


Acanthocheilonema  perstans,  Manson,  1891. 

Syn.  :   Filaria  perstans,  P.   Manson,  1891  ;   Filaria  sanguinis  hominis  var.  ininor, 

Manson,  1891. 

The  adult  female  Ac.  perstans  measures  70  to  80  mm.  in  length 
by  120  fi  to  140  fjL  in  breadth.  The  head  is  club-shaped  and  measures 
0*07  mm.   in   diameter.      The   vulva  opens  at  ob  to  I'o  mm.  from 


ACANTHOCHEILONEMA    PERSTANS 


415 


the  head.  The  tail  is  curved  and  presents  a  cuticular  thickening  which 
forms  two  triangular  appendages.  The  anus  opens  at  the  apex  of  a 
papilla  situated  in  the  concavity  of  the  curve  formed  by  the  tail  150  fju 
from  the  end.  The  diameter  of  the  tail  just  before  termination  is 
o'02  mm. 


Fig.  2gg.—Mf.  Persians  in  thick  film,  dried  and  stained  with  hsematoxylin 
4,  unshrunken  ;  5,  shrunken,      x  1,000.     (After  Fulleborn.) 


The  adult  male  measures  45  mm.  in  length  by  60  />t  to  80  /a  in 
breadth.  The  diameter  of  the  head  is  0*04  mm.  The  tail  is  much 
curved.  There  are  four  pairs  of  pre-anal  papillae  and  two  pairs  of 
post-anal  papillae.  Spicules  very  unequal  in  size.  Cloaca  121  /a  from 
the  tail  end.     At  the  tail  end  two  triangular  cuticular  appendages. 


4i6 


THE   ANIMAL   PARASITES   OF   MAN 


The  adalt  worms  inhabit  the  connective  tissue  at  the  base  of 
the  mesentery,  especially  in  the  region  of  the  pancreas,  abdominal 
aorta  and  suprarenals.  To  find  them  the  mesentery  should  be 
removed,  placed  in  a  2  per  cent,  solution  of  formalin,  and  then 
carefully  examined  at  leisure. 

Mikrofilaria  perstans. — 160  fiio  210  fi  by  ^  fi  to  6  /jl  broad.  Has  no 
sheath.  Cuticle  transversely  striated.  Tail  rounded  off,  not  pointed. 
Nerve  ring  at  34  fi.  Excretory  pore  49  fi,  genital  pore  125  fju  from 
head.  Smaller  larvae  90  fju  to  no  /a  by  4  yit  broad.  A  "  fang"  is  also 
described  on  the  head. 

M/.  Persians.  Mf.  demarquayi. 

(i)  Tail  stumpy.  (i)  Tail  pointed. 

(2)  Column  of  nuclei  extends  to  tip  of  (2)  Does  not  extend  to  tip. 

tail. 

Periodicity. — N  one. 
Life-history. — Unknown. 

Geographical  Distribution. — Very  common  in  many  parts  of  Africa  : 
Sierra  Leone,  Dahomey,  Northern  Nigeria,  Southern  Nigeria,  Came- 
roons,  Ivory  Coast,  Gold  Coast,  Old  Calabar,  Congo,  Uganda.  Absent 
from  Zululand,  Basutoland.  On  the  East  Coast  of  Africa  it  is  not 
found  in  the  towns  of  Zanzibar  and  Mombasa,  neither  is  it  found  in 
the  country  of  the  Masi,  nor  amongst  the  Kavirondo,  who  dw^ell 
along  the  north-east  shores  of  Lake  Victoria. 

In  South  America,  Ac.  perstans  is  very  common  amongst  the 
aboriginal  Indians  in  the  interior  of  British  Guiana.  However, 
it  is  not  found  in  Geoi-getown  and  in  New  Amsterdam,  neither  is 
it  found  in  the  cultivated  strip  of  coast  lying  between  these  two 
towns,  but  it  is  common  on  the  coast  farther  north  near  the  Vene- 
zuelan boundary,  where  the  forests  stretch  to  the  sea.  The  Waran 
Indians,  who  live  at  the  mouth  of  the  Waini  river,  harbour  this 
parasite.     It  is  absent  in  the  West  Indies. 

Topographically,  Ac.  perstans  is  found  only  in  areas  covered  by 
dense  forest  growth  and  abounding  in  swamps.  In  Kavirondo, 
where  the  forest  disappears  and  the  land  is  covered  with  scrub  and 
short  grass,  it  is  not  found  ;  likewise  it  is  not  found  on  tlie  grassy 
plains  of  the  highlands  of  British  East  Africa.  Towns  and  cultivated 
areas  are  free  from  it. 

Genus.  Dirofilaria.  Railliet  and  Henry,  191 1. 
Body  very  long,  thread-like,  cuticle  transversely  striated.  Mouth 
with  six  papillae.  Male  tail  spiral  with  voluminous  pre-anal  and  some 
large  post-anal  papillae  ;  spicules  unequal.  Vulva  near  the  anterior 
hundredth  of  body;  viviparous.  Parasitic  in  heart  or  blood-vessels 
and  subcutaneous  tissue. 


ONCHOCERCIN^ 


417 


DIrofilaria  magalhaesi,  R.  Blanchard,  1895. 


Fig.  300. — Dirojilaria 
magalhaesi :  posterior  ex- 
tremity. (After  V.  Lin- 
stow.) 


Syn.  :  Filaria  bancrofti^  v.   Linstovv,  1892  ;  Filaria  bancrofti^  P.  S.  de  Magalhaes, 

1892  inec  Cobbold,  1877). 

The  male  measures  83  mm.  in  length  by  0*28  to  0-40  mm.  in 
breadth.  The  anterior  extremity  is  rounded,  and  has  no  papillae  (?  6) ; 
the  posterior  extremity  exhibits  a  double  curve, 
with  four  pre-anal  and  four  post-anal  papillae  on 
each  side.  These  are  large  and  have  a  villous 
appearance.  The  mouth  is  round  and  unarmed, 
the  pharynx  measures  i  mm.  in  length,  is  cylin- 
dfical,  very  muscular,  and  its  hinder  part  is 
dilated.  The  anus  is  situated  o'li  mm.  in  fron 
of  the  hind  end.  There  are  probably  two 
unequal  spicules;  one  only,  however,  is  known 
— apparently  the  shorter  one — the  length  of 
which  is  given  as  o'ly  to  0*23  mm. 

The  female  measures  155  mm.  in  length 
and  0*6  to  o*8  mm.  in  breadth  ;  the  rings  of 
the  cuticle  are  0*005  nam.  apart  (in  the  male 
0-003  mm.  apart)  ;  the  anterior  extremity  is 
slightly  thickened  and  club-like,  the  posterior 
extremity  is  slender,  and  terminates  obtusely ; 
the  lateral  line  is  o'i27  mm.  in  breadth  (that  of 

the  male  0*007  to  o*oo8  mm.);  the  anus  opens  o'i3  mm.  in  front  of 
the  hind  end,  the  vulva  is  2*5  mm.  distant  from  the  mouth,  the  ovaries 
are  two  much  convoluted  tubes.     The  eggs  measure  38  /><,  by  1 1  /i,. 

This  species  was  first  discovered  at  a  post-mertem^  in  the  left  ventricle,  by  J.  P. 
Figueira  de  Saboia  in  Rio  de  Janeiro,  and  has  been  described  by  P.  S.  de  Magalhaes. 

D.  immitis  occurs  in  the  right  ventricle  of  the  heart  of  the  dog  in  Europe  and 
the  Tropics. 

D.  repens  is  also  a  common  subcutaneous  Nematode  in  dogs  in  Annam. 

Sub-family.     Onchocercinae,  Leiper,  191 1. 
Cuticle  with  spiral  thickenings. 

Genus.     Onchocerca,  Diesing,  1841. 

Male  with  four  pre-anal  papillae.  Female  with  vulva  situated 
anteriorly. 

Onchocerca  volvulus,  R.  Leuckart,  1893. 

Syn.  :  Filaria  volvulus^  R.  Leuckart,  1893. 
The  adult  male  measures  30  to  35  mm.  in  length  by  0*14  mm.  in 
breadth.     The  body  is  white,  filiform,  attenuated  at  both  ends.     The 
head  is  rounded   and    has   a    diameter    of   0*048  mm. 


The   cuticle 


4l8  THE   ANIMAL   PARASITES   OF    MAN 

is  distinctly  transversely  striated.  The  mouth  is  unarmed.  The 
alimentary  canal  is  straight,  the  anus  opening  q-o;  mm.  from  the  tip 
of  the  tail.  The  tail  is  strongly  curved  and  somewhat  flattened  on 
the  concave  surface.  There  are  three  papillae,  one  large  and  two 
small,  on  each  side  of  the  cloaca  and  one  large  and  two  post-anal 
small  papillae.     Two  curved  spicules,  0*166  and  0*08  mm.  respectively. 

The  adult  female  is  of  uncertain  length,  but  much  longer  than 
the  male,  probably  about  10  to  12  cm.  The  head  is  rounded  and 
truncated  ;  it  measures  0*065  mm.  in  diameter.  The  tail  is  curved. 
The  vulva  opens  0*55  mm.  from  the  head.  The  hand-like  cuticular 
thickenings  are  well  marked.  Eggs  ovoid  with  a  prolongation  at 
each  pole  'Mike  an  orange  wrapped  in  tissue  paper."  The  larva 
measures  about  300  yu,  by  7  /a  to  S  fi;  it  has  no  "sheath."  The  body 
tapers  from  about  the  last  fifth  of  its  length,  and  terminates  in  a 
sharply  pointed  tail.  At  about  the  anterior  fifth  of  the  body  there 
is  a  V  spot. 

0.  volvulus  is  found  in  peculiar  subcutaneous  tumours,  the  size  of 
a  pea  to  that  of  a  pigeon's  Qgg.  The  same  patient  may  present  one 
or  several  of  these  tumours.  The  regions  of  the  body  most  fre- 
quently affected  are  those  in  which  the  peripheral  lymphatics  con- 
verge. Thus  they  are  usually  found  in  the  axilla,  in  the  popliteal 
space,  about  the  elbow,  in  the  sub-occipital  region  and  in  the  inter- 
costal spaces.  The  tumours  are  never  adherent  to  the  surrounding 
structures,  and  can  be  easily  enucleated.  They  are  formed  of  a  dense 
connective  tissue  wall  and  internally  a  looser  fibrous  meshwork. 
This  is  traversed  by  a  series  of  canals  in  which  the  worms  lie,  but 
they  are  also  partly  embedded  in  the  denser  wall.  The  canals 
apparently  dilate  into  Cavities  filled  with  slimy  pus-like  fluid  con- 
sisting largely  of  larvae.  According  to  Brumpt  the  posterior  extremity 
of  the  male,  and  the  anterior  extremity  of  the  female  with  its  vaginal 
opening,  are  free  in  one  of  the  spaces  for  the  purpose  of  copulation 
and  parturition.  If  a  tumour  be  cut  into  and  placed  in  salt  solution, 
Rodenwaldt  states  that  the  undamaged  males  wander  out  into  the 
solution. 

The  formation  of  the  tumours  is  elucidated  by  Labadie-Lagrave 
and  Deguy's  case.  The  authors  found  an  immature  female  Onchocerca 
volvulus  in  a  lymphatic  vessel  partly  obstructed  by  an  infiltration  of 
fibrin  and  leucocytes.  It  appears,  therefore,  that  the  presence  of  the 
parasites  within  the  lymphatics  gives  rise  to  an  inflammatory  process, 
and  that  the  consequent  fibrinous  deposit  envelops  the  parasites, 
obliterates  the  lumen  of  the  vessel,  and  ultimately  isolates  the  affected 
tract.  At  any  rate,  in  young  tumours  the  worms  appear  to  lie  in 
a  structureless  substance  permeated  by  leucocytes  in  which  connective 
tissue  is  gradually  organized  from  the  periphery,  thus  isolating  the 
worms. 


TRICHINELLID^ 


419 


In  cases  of  infection  with  0.  volvulus  larvae  have  been  found  by 
Onizilleau,  Fiilleborn,  and  Simon  in  lymph  glands,  and  in  the  finger 
blood  if  considerable  pressure  is  used  so  as  to  squeeze  lymph  out 
of  the  tissues.  They  are  sheathlesSj  and  the  following  are  the 
dimensions  in  ordinary  dried  films:  Length,  274 /^ ;  nerve  ring, 
23-7  per  cent.  ;  Gi  cell,  6()'6  per  cent.  ;  end  of  last  tail  cell,  96*3  per 
cent.  The  dimensions  of  larvae  of  0.  volvidiis  taken  from  the  uterus 
and  prepared  in  the  same  way  are:  Length,  224*5 yu,;  nerve  ring, 
24-3  per  cent. ;  Gi  cell,  68*9  per  cent.  ;  end  of  the  last  tail  cell, 
95*5  pci'  cent.  In  all  probability  the  larvae  in  the  glands  and  blood 
are  those  of  0.  volvulus. 

According  to  the  natives,  the  tumours  may  last  indefinitely  and 
never  ulcerate.  Some  old  patients  told  Brumpt  that  their  tumours 
had  been  present  since  childhood.  Probably  Onchocerca  volvulus, 
like  some  other  Filariidce,  may  live  for  many  years. 

0.  volvulus  occurs  in  various  parts  of  West  Africa  :  Gold  Coast, 
Sierra  Leone,  Dahomey,  Lagos,  Cameroons.  Brumpt,  on  the  banks 
of  the  Welle  between  Dongon  and  M'Binia  (Belgian  Congo),  found 
about  5  per  cent,  of  the  riverine  population  affected. 

Family.     Trichinellidae,  Stiles  and  Crane,  1910. 
Sub-family.     Trichurinae,  Ransom,  191 1. 

Male  with  a  single  long  spicule,  with  sleeve-like  sheath.  One 
ovary.  Eggs  with  an  opening  at  each  pole  closed  by  a  plug-like 
operculum.  Eggs  hatch  on  being  swallowed  by  a  new  host.  Genera  : 
Trichuris,  Capillaria. 

Genus.     Trichuris,  Roderer  and  Wagler,  1761. 

Syn. :     Trichocephalus,  Goeze,  1782  {7iec  Trichiurus,  L.,  1758);    Mastigodes^ 

Zeder,   1803. 

The  anterior  part  of  the  body  is  very  long  and  thread-like  :  the  posterior,  much 
shorter  part,  is  thicker,  rounded  posteriorly,  and  the  anus  is  terminal.  The  males 
have  the  posterior  extremity  spirally  rolled  ;  the  vulva  is  situated  at  the  commence- 
ment of  the  posterior  part  of  the  body.  The  Trichocephali  live  in  the  large  intestine 
of  mammals,  the  caecum  by  predilection  ;  their  development  is  direct,  infection  occurs 
through  the  ingestion  of  embryo-containing  eggs. 

Trichuris  trichiura,  Linnaeus,  1761. 

Syn.:     Trichocephaliis    trichiurus.,    L.,    1771  ;    Ascaris   trichiura,    L.,    1771  ; 
Trichocephalus  hominis,  Schrank,  1788;   Trichocephalus  dispar.,  Rud.,  1801. 

The  male  measures  40  to  45  mm.  in  length,  the  spicule  is 
2*5  mm.  long,  its  retractile  sheath  is  beset  with  spines.  The  female 
measures  45    to    50   mm.   in    length,    of   which   two-fifths   appertain 

27 


4^0 


THE   ANIMAL   PARASITES   OF   MAN 


to  the  posterior  part  of  the  body.  The  ova  are  barrel-shaped  and 
have  a  thick  brownish  shell  which  is  perforated  at  the  poles.  Each 
opening  is  closed  by  a  light-coloured  plug.  The  eggs  measure 
50  fi  to  54  yLt  in  length  and  23  fi  in  breadth ;  they  are  deposited 
before  segmentation.  Trichuris  trichinra  usually  lives  in  the  caecum 
of  man,  and  is  also  occasionally  found  in  the  vermiform  appendix 
and  in  the  colon,  exceptionally  also  in  the  small  intestine  ;  usually 
only  a  few  specimens  are  present,  and  these  do  not  cause  any  parti- 
cular disturbance,  although,  as  Askanazy  found,  they  feed  on  blood ; 
in  other  cases  cerebral  symptoms  of  more  or  less  severity  are  observed 
when  Trichocephali  are  present  in  large  numbers.  At  posi-nwrtems 
performed  soon  after  death  the  filiform  anterior  extremity  of  the  worm 

is  frequently  found  embedded  in  the 
mucous  membrane  (Askanazy). 

The  whip  worm  is  one  of  the  most  common 
parasites  of  man  and  appears  to  be  distributed 
over  the  entire  surface  of  the  globe  ;  it  is,  how- 
ever, more  frequent  in  the  warmer  regions.  It 
is  found  in  persons  of  both  sexes  and  all  ages 
with  the  exception  of  infants.  In  autopsies  it 
is  found  in  the  following  numbers  :  In  Dresden 
in  2*5  per  cent.,  in  Erlangen  in  ii'i  per  cent., 
in  Kiel  in  31 '8  per  cent.,  in  Munich  in  9-3  per 
cent.,  in  Petrograd  in  o'i8  per  cent.,  in 
Gottingen  in  46"i  per  cent.,  in  Basle  in  237  per 
cent.,  in  Greenwich  in  68  per  cent.,  in  Dublin 
in  89  per  cent.,  in  Paris  in  about  50  per  cent., 
and  in  Southern  Italy  in  almost  100  per  cent. 
On  examining  the  fasces  the  eggs  of  the  whip 
worm  were  found  as  follows  :  In  Munich  in  8*26 
per  cent.,  in  Kiel  in  45*2  per  cent.,  in  Greifswald 
in  45  per  cent.,  in  North  Holland  in  7  per  cent., 
in  Novgorod  in  26*4  per  cent.,  in  Petrograd 
in  5  per  cent.,  in  Moscow  in  5*3  per  cent. 


Fig.  301. — Trichufis  trichiui-a  :  on 
the  left,  male  ;  on  the  right,  female  with 
the  anterior  extremity  embedded  in  the 
mucous  membrane  of  the  intestine ; 
below,  egg. 


The  development  of  the  eggs  is  completed  in  water  or  in  moist 
soil,  and  occupies  a  longer  or  shorter  time  according  to  the  season  ; 
the  eggs  possess  great  powers  of  resistance,  as  do  the  larvae,  which, 
according  to  Davaine,  may  remain  as  long  as  five  years  in  the  egg- 
shell without  losing  their  vitality.  Leuckart  proved  by  experiment 
that  direct  infection  with  Trichuris  ovis  {Ovis  aries)  and  T.  crenata 
(Siis  scrofa  dom.)  was  produced  by  embryo-containing  eggs  ;  Railliet 
obtained  the  same  results  with  T.  depressiusciila  of  dogs,  and  Grassi 
subsequently,  by  rrieans  of  two  experiments,  demonstrated  the  direct 
development  of  Trichuris  trichinra.  In  one  case  embryo-containing 
eggs  were  swallowed  on  June  27,  1884,  and  on  July  24  the  ova  of 
Trichocephali  were  found  in  the  faeces  for  the  fii-st  time. 


TRICHINELLIN^  42I 

Trichuris  trichiiira  is  found  not  only  in  man,  but  also  in  various 
monkeys  {T.  palceformis,  Rud.),  as  well  as  in  lemurs  (T.  lemur  is,  Rud.)- 

Other  species  are  T.  crenata  in  pig  ;  T.  ovis  in  cattle,  sheep,  goat,  and 
pig  (?)  ;  T.  depressiusciila  in  dog  ;  T.  campanula  in  cat ;  T.  unguiculata 
in  rabbit  and  hare  ;  T.  cameli  in  camel;  T.  discolor  in  humped  cattle  ; 
T.  nodosus  in  mouse ;  T.  alcocki  in  the  thamin  (India)  ;  T.  globidosa  in 
camel  ;  T.  giraffce  in  giraffe. 

Sub-family.     Trichinellinae,  Ransom,  191 1. 

Male  without  spicule  ;  females  ovoviviparous.  Larvae  penetrate 
muscles  of  host  and  become  encysted.     Genus  :  Trichinella. 

Genus.     Trichinella,  Railliet,  1895. 

Syn.  :   Trichina^  Owen,  1835  (^^^  Meigen,  1830). 
Very  small  Trichinellincs,  the  males  of  which  have  two  conical  appendages  at  the 
•caudal  extremity  ;    the  vulva  is   situated  at  the  border  of  the  anterior  fifth  of  the 
body.     There  is  only  one  species. 

Trichinella  spiralis,  Owen,  1835. 

Syn. :   Trichina  spiralis^  Owen,  1835. 

The  male  measures  i"4  to  i'6  mm.  in  length  and  0*04  mm.  in 
diameter.  The  anterior  part  of  the  body  is  narrowed,  the  orifice  of 
the  cloaca  is  terminal  and  lies  between  the  two  caudal  appendages  ; 
internal  to  these  are  two  pairs  of  papillae,  dorsal  one  behind  the  other. 
.The  cloaca  is  evertible  for  copulation.  The  females  measure  3  to 
4  mm.  in  length  and  o'o6  mm.  in  diameter  ;  anus  terminal. 

Trichinella  spiralis  in  its  adult  stage  inhabits  the  small  intestine 
of  man,  pig,  wild  boar,  rat.  The  young  do  not  leave  the  body  of  the 
host  but  become  encysted  in  the  muscles.  Experimentally  it  develops 
in  the  black  rat  (Mus  rattus),  the  sewer  rat  {M.  decumamis),  the 
■domestic  pig  (Sus  scrofa  dom.),  the  wild  boar  {Siis  scrofa  ferox),  the 
domestic  dog  {Canis  familiaris) ,  the  fox  (C.  vulpes)  the  badger  (Meles 
iaxiis),  the  polecat  (Putorius  fcetidits),  the  marten  (Mustela  /oina),  the 
raccoon  (Procyon  lotor),  the  hippopotamus  and  the  cat,  and  many 
other  mammals  (rodents  and  carnivora) ;  Trichinellae  have  been  arti- 
ficially introduced,  by  administering  the  encysted  stage,  into  the  dog, 
the  mole  {Talpa  europcea),  the  mouse  (Mus  musculiis),  the  hare  {Lepus 
timidus),  the  rabbit  (L.  cuniculiis),  the  hedgehog  {Eriiiaceus  europcBus), 
the  marmot  {Cricetus  vulgaris),  the  vole,  the  dormouse,  the  sheep,  the 
calf,  the  horse,  etc.  Human  beings  and  the  pig,  rat,  mouse,  guinea- 
pig  and  rabbit  are  most  easily  infected ;  less  easily  the  sheep,  calf  and 
horse  ;  with  difficulty  the  cat,  dog  and  badger.  Trichinella  can  also 
be  reared  in  birds  (fowl,  pigeon  and  duck),  but   the  young  do  not 


422 


THE   ANIMAL   PARASITES   OF   MAN 


encyst  in  the  muscular  system,  but  are  expelled  with  the  faeces.     By 
cold-blooded   animals   as   well  as  by  insects    (Calliphora  vomitaria) 


Fig.  302. — Trichinella  spiralis.  ?,  mature  female  :  E^  embryos;  V^  vulva;  Ov,  ovary. 
i  ,  mature  male  :  7",  testes,  c,  newly  born  larva,  d.,  larva  in  the  muscles.  <?.,  encapsuled  larva 
in  the  muscles.     Magnified.     (After  Glaus.) 

encysted  Trichinellae  are  evacuated  without  undergoing  any  change, 
but  they  will  still  develop  if  subsequently  ingested,  say,  by  rabbits. 
According  to  Gujon,  however,  Trichinella  can  develop  in  salamanders. 


TRICHINELLA   SPIRALIS  423 

because  he  -has  found  Trichinella  of  the  muscles  in  these  animals 
after  they  had  been  fed  on  encysted  specimens.  A  high  temperature 
(30°  C.)  must  be  provided  in  which  to  keep  the  experimental  animals 
to  ensure  the  success  of  the  infection. 

History. — Encapsuled  Trichinella  had  been  observed  in  London  by  Peacock 
(1828)  and  by  J.  Hilton  (1833)  in  the  muscular  system  of  man  ;  soon  after 
(1835),  Paget  found  them  in  London  in  an  Italian  who  had  died  of  tuberculosis, 
and  recognized  them  to  be  encysted  entozoa,  which  R.  Owen  described  as  Trichifta 
spiralis.  Soon  after,  some  further  observations  were  reported  on  the  occurrence  of 
encysted  Trichinellae,  in  man^  in  England,  Berlin,  Heidelberg,  Denmark,  North 
America  ;  they  were  also  found  in  the  pig  (Leidy,  Philadelphia)  and  the  cat  (Herbst, 
Gottingen,  and  Gurlt,  Berlin).  Herbst  even  succeeded  in  infecting  a  badger  with 
encysted  Trichinellae,  and  subsequently  infected  two  dogs  with  the  flesh  of  this 
badger  (1850).  In  1855  R.  Leuckart  (Giessen)  also  commenced  feeding  experiments, 
and,  like  Kiichenmeister  and  Virchow  (1859),  first  went  on  the  wrong  track  because 
it  was  believed  at  that  time  that  Trichinellae  were  the  larvas  either  of  Trichocephalus 
or  Strongylus.  Nevertheless,  these  experiments  yielded  some  important  results  ; 
they  showed  that  Trichinellae  become  adult  in  the  intestine  within  a  few  days,  and 
that  the  females  are  viviparous  (Leuckart).  Until  that  time  Trichinellae  had  been 
regarded  as  fairly  harmless  guests  of  man,  but  opinions  soon  changed  when  Zenker 
in  Dresden  (January,  i860),  in  performing  the  autopsy  of  a  girl,  aged  10,  who  had 
entered  the  hospital  with  typhoid  symptoms  and  there  died,  found  Trichinellae  (not 
yet  encysted)  in  the  muscles ;  the  intestinal  lesions  characteristic  of  typhoid  were 
lacking,  but  numerous  adult  Trichinellae  were  found  in  the  intestine.  Inquiries 
elicited  the  fact  that  at  about  Christmas  time  the  girl  had  been  taken  ill  after  eating 
pork,  and  at  the  same  time  the  butcher  from  whom  the  meat  was  brought  as  well 
as  several  of  his  customers  fell  sick  :  the  pickled  pieces  of  the  same  meat  were  full 
of  Trichinellae.  In  the  face  of  this  information  it  was  not  difficult  to  ascertain  the 
cause  of  the  disease  and  the  manner  of  infection  in  Zenker's  case,  and  it  was  not 
long  before  Leuckart,  Virchow  and  Zenker  were  able  by  renewed  experiments  to 
demonstrate  the  cycle  of  development  of  Trichinella  spiralis.  Similar  investigations 
followed  by  Glaus  in  Wiirzburg,  Davaine  in  Paris,  Fuchs  and  Pagenstecher  in 
Heidelberg,  etc. 

Hardly  had  Zenker's  case  been  published  than  numerous  observations  on 
trichinosis  in  man  appeared,  some  referring  to  isolated  cases,  others  to  small  or 
great  epidemics,  and  nearly  all  from  North  Germany.  The  worst  epidemic  was 
that  of  Hadersleben  (1865),  in  which  place,  numbering  hardly  2,000  inhabitants, 
337  persons  were  taken  ill  within  a  short  time,  and  of  these  loi  died.  The  source 
of  infection  proved  to  be  a  single  pig,  the  flesh  of  which  had  been  mixed  with  that 
of  three  other  pigs  :  200  of  the  badly  infected  persoi^s  had  exclusively  eaten  raw 
pork. 

Moreover,  it  soon  became  clear  that  epidemics  of  trichinosis  had  been  observed 
in  Germany  prior  to  i860,  but  that  their  nature  had  not  been  recognized,  although 
in  a  few  cases  Trichinellae  had  been  found  in  the  muscles  of  those  who  had 
succumbed. 

History  of  the  Development  of    Trichinella  spiralis. 

Shortly  after  their  introduction  into  the  intestine  of  experimental 
animals  the  encysted  Trichinellae  escape  from  their  capsules,  which 
are  destroyed  by  the  gastric  juices,  and  they  then  enter  the  duodenum 


424  THE   ANIMAL   PARASITES   OF   MAN 

and  jejunum,  where  they  become  adult.  During  this  period  they  do 
not  grow  much,  the  males  from  o*8  to  I'o  to  i'2  to  1*5  mm.;  the 
females  to  1*5  to  i'8  mm.  Soon  after  copulation,  which  takes  place 
in  the  course  of  two  days,  the  males  die  ;  the  females,  w^hich  during 
the  following  days  attain  a  length  of  3  to  3*5  mm.,  either  bore  more 
or  less  deeply  into  the  villi  or,  by  means  of  Lieberkuhn's  glands,  into 
the  mucous  membrane  (Askanazy,  Cerfontaine,  Geisse),  and  thus 
usually  attain  the  lymph  spaces.  A  few  also  pierce  the  intestinal  wall 
and  are  then  found  in  the  mesentery  and  glands.  The  females  deposit 
their  young,  the  number  of  which,  according  to  Leuckart,  averages  at 
least  1,500,  in  the  lymph  spaces  ;  the  newly  born  larvae  measure  90//,  to 
100  ^  in  length,  6  //<  in  diameter,  and  they  do  not  appear  to  increase  in 
size  during  their  migrations.  The  migrations  are  mostly  passive,  that 
is  to  say,  the  larvae  are  carried  along  mainly  by  the  lymph  stream  to 
the  heart,  but  sometimes  they  are  active,  as  may  be  inferred  from  the 
fact  that  young  Trichinellae  are  found  in  various  parts  of  the  intes- 
tinal w^all  beyond  the  chyle  and  lymph  spaces,  as  well  as  in  abun- 
dance in  the  abdominal  cavity.  Trichinellae  occur  in  the  heart's 
blood  of  artificially  infected  animals  seven  to  twenty-three  days 
after  infection.  If  scanty,  dilute  the  blood  w^ith  about  ten  times  the 
amount  of  3  per  cent,  acetic  acid  and  centrifugalize. 

The  young  brood  is  distributed  from  the  heart  throughout  the 
entire  body,  but  the  conditions  necessary  to  its  further  development 
are  found  only  in  striated  muscle  ;  the  young  Nematodes  penetrate 
the  capillaries,  attain  the  intramuscular  connective  tissue  and  then 
invade  the  fibres  (Virchow,  Leuckart,  Graham^).  On  the  ninth  or 
tenth  day  after  infection  the  first  Trichinellae  have  reached  their 
destination  ;  but  further  invasions  are  constantly  taking  place  because 
the  intestinal  Trichinellae  live  from  five  to  seven  weeks,  and  continue 
to  produce  their  young. 

Symptoms. — (i)  Period  of  invasion  :  Gastro-intestinal  symptoms — 
nausea,  vomiting,  watery  diarrhoea,  colic.  Muscular  pains  may  occur 
even  at  this  period.  Recurrent  abdominal  pains  about  the  eighth  day, 
a  temporary  oedema.     Embryos  are  abundant  in  the  serous  cavities. 

(2)  Period  of  dissemination  :  Second  week.  Myositis,  variable  in 
amount,  is  the  predominant  symptom.  The  biceps  and  calf  may  be 
hard  and  tender.  Mastication,  speech,  respiration,  etc.,  may  be 
difficult  and  painful.  Dyspnoea  may  be  intense.  Temperature 
104°  to   105°  F. 


'  TrichinellK  that  are  unable  to  penetrate  into  muscular  fibres  invariably  die,  no  matter 
where  else  they  settle  ;  their  occurrence  in  the  adipose  tissue  is  disputed,  but  is  still  possibly- 
correct,  as  bundles  of  muscles  are  present  in  the  fat  of  bacon.  The  Trichinellae  do  not  settle 
in  heart  muscle,  although  they  may  reach  it  in  cases  of  heavy  infection  ;  they  then  die  or 
wander  into  the  pericardium,  and  eventually  into  the  heart  cavities. 


TRICHINELLA    SPIRALIS 


425 


(3)  Period  of  encystment :  Symptoms  of  marked  cachexia.  Third 
week  :  Second  period  of  oedema,  especially  of  face.  Delirium, 
somnolence,  lung  affections.  Death  or  gradual  subsidence  of 
symptoms    in    mild    cases. 

Eosinophilia  (50  per  cent,  or  more)  is  present. 

In  consequence  of  the  new  batches  of  young  produced  during  several  weeks, 
the  above-mentioned  symptonis  of  disease  are  often  considerably  aggravated  ;  the 
fever  increases,  delirium  may  arise,  and  infiltration  of  the  lungs,  fatty  degeneration 
of  the  liver  and  inflammation  of  the  kidneys  may  ensue;  the  initial  slight  oedema 


Fig.  303. — A.,  isolated  muscular  fibre  of  a  rat,  invaded  by  Trichinella.  510/1.  B., 
section  through  the  muscle  of  a  rat;  the  infected  fibre  has  lost  its  transverse  stiiation  ;  its 
nuclei  are  enlarged  and  multiplied.  310/1.  C,  portion  of  a  Trichinella  capsule,  at  the  pole 
of  which  connective  tissue  cells  are  penetrating  tlie  thickened  sarcolemma.  (After  Hertwig- 
Graham.) 

may  extend,  the  strength  dwindles,  and  in  many  cases  the  patients  succumb  to  the 
trichinosis.  In  severe  cases  improvement  of  the  condition  is  only  apt  to  occur 
in  the  fourth  or  fifth  week  ;  the  convalescence  is  always  protracted. 

The  muscular  fibres  attacked  degenerate,  the  transverse  striation 
at  first  disappearing  ;  the  fibres  then  assume  a  granular  appearance, 
the  nuclei  multiply  and  become  enlarged,  and  are  surrounded  by  an 
area  of  granular  material,  which  stains  more  deeply  than  the  remaining 
contents  of  the  sarcolemma.  Two  or  three  weeks  after  infection,  the 
spirally  rolled-up  Trichinelhie  have  grown  to  o*8  to  I'o  mm.,  and  in 
their  vicinity  the  muscular  fibre  is  swollen,  spindle-shaped,  and  the 


426  THE   ANIMAL  PARASITES   OF   MAN 

sarcolemma  is  glassy  and  thickened.  The  inflammation  also  extends 
to  contiguous  fibres,  especially  to  the  intramuscular  tissue,  which 
proliferates  greatly,  especially  in  the  vicinity  of  the  degenerated  fibres. 
While  the  latter  become  more  and  more  absorbed,  the  capsule  is 
formed  by  the  inflamed  connective  tissue,  which,  penetrating  into  the 
glassy  and  thickened  sarcolemma  from  the  poles  of  the  spindle,  forms 
the  cystic  membrane.  According  to  other  authorities,  the  larvae  settle 
in  the  ////^rmuscular  connective  tissue  which  forms  the  cyst  and  not 
in  the  muscular  fibres  within  the  sarcolemma.  The  cysts  are  lemon- 
shaped  and  usually  lie  with  their  longitudinal  axis  in  the  direction  of 
the  muscular  fibres  ;  on  an  average  they  measure  400  ^  in  length  by 
250  /x  in  breadth. 

Later  on  fat  cells  appear  at  their  poles,  and  after  about  six  or 
nine  months  they  commence  to  calcify,  the  process  starting  at  the 


Fig.    305. — Various  phases  of  the  calcification  of 
,  Trichinella  of  the  muscles,  which  starts  at  the  poles 

Fig.  304.-Calcified  Trichinella  in  the    ^^  ^^^  ^'P'"^^" 
muscular  system  of  a  pig ;  the  capsules  are 
not  calcified.     (After  Ostertag. ) 

poles  (fig.  305).  Finally,  sometimes  after  the  lapse  of  years,  the 
captive  Trichinellae  themselves  become  calcified. 

According  to  experience,  Trichinellae  are  not  evenly  distributed 
in  the  muscular  system  of  pigs  ;  the  diaphragm,  the  muscles  of  the 
larynx,  tongue,  abdomen  and  intercostal  spaces  are  their  favourite 
positions  ;  this  predilection  for  the  respiratory  muscles  is  explained 
by  their  regular  contractions,  owing  to  which  regular  narrowings 
of  the  capillaries  take  place,  thus  favouring  the  settling  of  the  cir- 
culating Trichinellae.  The  same  circumstance  probably  explains  the 
frequency  of  the  parasites  in  the  tongue. 

Possibly  also  the  Trichinellae  that  bore  direct  through  the  intestine 
may,  from  the  abdominal  cavity,  penetrate  the  muscles  in  the  vicinity. 
Frequently  also  encysted  Trichinellae  are  found  in  remarkable  numbers 


TRICHINELLA   SPIRALIS  427 

in  the  vicinity  of  the  points  of  insertion  of  the  tendons,  this  pro- 
cHvity  being  probably  connected  with  the  fact  that  the  Trichinellae 
first  of  all  wander  into  the  muscular  fibres  and  find  a  natural  barrier 
at  the  points  of  insertion  of  the  tendons. 

The  Trichinellae,  in  their  encysted  condition,  may  remain  alive 
and  capable  of  development  for  many  years — in  the  pig  eleven  years 
and  in  man  as  much  as  twenty-five  to  thirty-one  years.  Encystment, 
however,  is  not  a  necessary  condition  for  the  development  of  the 
brood,  that  is  to  say,  Trichinellae  which  reach  the  gut  of  suitable 
animals  become  sexually  mature  and  multiply  provided  that  they  have 
developed  so  far  as  to  possess  a  rudimentary  genital  spot,  which 
occurs  when  the  body  is  0*5  to  075  mm.  long,  but  all  the  same 
a  great  part  of  non-encapsuled  Trichinae  perish  on  their  passage 
through  the  stomach. 

The  black  rat  (Mus  rattus),  and  more  particularly  the  sewer  rat 
(Mas  deaimanns^),  are  the  normal  hosts  of  Trichinella  spiralis.  These 
animals,  especially  the  last-named  species,  infect  themselves  very 
easily,  as  they  are  cannibalistic,  and  they  also  transmit  trichinosis 
to  other  species  by  which  they  are  devoured,  such  as  pigs,  dogs, 
cats,  foxes,  bears  and  martens.  Rats  are  infected  also  by  the  in- 
gestion of  faecal  matter  from  infected  animals  which  contains 
trichinae  (Hoyberg).  Man  becomes  infected  with  Trichinella  by 
eating  the  flesh,  insufficiently  cooked,  of  infected  pigs,  also,  but 
more  rarely,  by  eating  the  infected  flesh  of  wild  boars,  dogs,  cats, 
bears  and  foxes. 

The  infection  of  pigs  may  likewise  take  place  by  their  having  access  to  the  offal 
of  trichinous  pigs,  or  being  actually  fed  on  it.  These  are,  however,  exceptions, 
which,  as  a  matter  of  course,  are  of  great  importance  in  certain  places.  As 
a  matter  of  fact,  the  rats  examined  for  Trichinella  were  always  found  to  be  severely 
infected.  Thus  Billings,  in  the  knackers'  yard  at  Boston,  found  that  76  per  cent, 
of  the  rats  were  infected,  and  in  an  export  slaughterhouse  100  per  cent,  were  found 
to  harbour  the  parasite  ;  in  the  city  of  Boston  10  per  cent,  of  the  rats  had  trichinosis. 
Heller  found  that  of  704  rats,  from  twenty-nine  different  places  in  Saxony,  Bavaria, 
Wiirtemberg  and  Austria,  8*3  per  cent,  were  infected  with  Trichinellce ;  of  the  rats 
caught  in  the  knackers'"  yards,  22' i  were  diseased;  of  those  taken  in  slaughter- 
houses, 2'3  were  infected,  and  of  rats  from  other  localities  only  0-3  per  cent, 
harboured  the  parasite.  Leisering  found  almost  the  same  figures,  but  in  rats  from 
slaughterhouses  5*3  per  cent  were  infected. 

The  geographical  distribution  of  T.  spiralis  does  not  correspond 
with    the   occurrence    of   trichinosis    in    man  ;    local    customs    are 

'  It  is  siill  a  matter  of  dispute  and  can  hardly  be  definitely  settled  whether  Trichinellse  were 
brought  to  Europe  by  the  sewer  rats  which  invaded  Europe  at  the  end  of  the  eighteenth 
century,  or  whether  they  were  imported  with  the  Chinese  pig  in  1820  or  1830,  when  it  was 
introduced  into  England  and  Germany  to  cross  with  the  native  breeds,  or  whether  finally 
Trichinellae  are  also  indigenous  to  Europe. 


428 


THE   ANIMAL   PARASITES   OF   MAN 


an  important  factor ;  for  instance,  the  custom  of  eating  pork  in 
a  condition  that  does  not  affect  the  Hfe  of  the  enclosed  trichinella. 
In  places  where  such  customs  do  not  prevail,  epidemics  do  not 
occur — at  the  most  there  are  isolated  cases  of  the  disease,  although 
there  be  a  great  number  of  infected  pigs.  The  following  conditions 
prevail  in  North  America :  In  Boston,  Billings  found  that  4  to  57  per 
cent,  of  the  pigs  examined  were  trichinous  ;  Belfield  and  Atwood 
found  that  8  per  cent,  were  infected  in  Chicago  ;  Salmon  found 
on  an  average  that  27  per  cent,  were  infected  (but  at  various  places- 
the  percentage  fluctuated  between  0*28  to  16*3  per  cent.),  yet  epidemics 
of  trichinosis  hardly  ever  occur  in  North  America,  and  only  isolated 
cases  of  the  disease  are  met  with  in  German  immigrants,  who  keep 
to  their  native  customs. 

This  report,  according  to  the  researches  of  H.  U.  Williams,  must  be  consider- 
ably modified.  This  author  has  examined  the  muscular  system  of  human  cadavers 
according  to  the  method  employed  by  inspectors  of  meat  for  pigs.  The  investiga- 
tions were  conducted  in  the  Pathological  Institute  of  the  University  of  Buffalo,  and 
the  observer  has  examined  50:5  bodies  since  1894,  of  which  ^7  (=  y2>4  P^r  cent.) 
were  invaded  by  Trichinella.  The  cases,  according  to  the  nationality,  are  divided 
as  follows  : — 


Trichinella 

Examined 

Percentage  of 

positive  results 

Absent 

Present 

Americans  :  — 

{a)  Whites           

207 

201 

6 

2-89 

(6)  Negroes         

70 

65 

5 

7-14 

British  and  Irish     

62 

57 

5 

8 -06 

Canadians 

12 

10 

2 

16-66 

Germans      

49 

43 

6 

12-24 

Italians         

12 

10 

2 

16-66 

Other  nationalities 

27 

27 

0 

0 

Nationality  unknown 

66 

65 

I 

I-5I 

Total 

505 

478 

27 

5'34 

It  is  worthy  of  remark  that  half  of  all  the  positive  cases  were  mental  patients^ 
who  were  found  to  be  affected  with  Trichinella  to  well-nigh  12  per  cent.  Trichinosis 
was  not,  however,"the  cause  of  death  in  any  case.  Very  frequently  the  Trichinella^ 
were  found  calcified  and  dead. 


Conditions  are  similar  in  most  countries  of  Europe,  where,  of 
course,  the  number  of  infected  pigs  is  considerably  smaller,  but 
the  disease  depends  less  on  this  than  on  the  way  in  which  the 
pork  is  prepared. 

Cases  of  trichinosis  have  been  known  to  occur  in  nearly  all 
the  countries  of  Europe;  further,  in  Egypt,  Algeria,  East  Africa. 
Syria,     India,     Australia,    and     America.       North     Germany,    more 


TRICHINELLA   SPIRALIS  429 

especially  the  Saxe-Thiiringian  states,  is  the  classical  land  for 
epidemics  of  trichinosis ;  the  mortality  varies,  but  it  may  be 
very  high.^ 

Prophylaxis. — The  grave  nature  of  the  disease  and  the  comparatively  high 
mortality  relating  to  trichinosis  led  the  authorities  to  adopt  certain  preventive 
measures,  which  are  the  more  necessary  as  national  customs  cannot  be  altered  in 
a  short  time.  As  the  usual  process  of  pickling  and  smoking,  even  when  long 
continued,  does  not  certainly  ensure  the  death  of  the  Trichinellse  contained  in  the 
meat,  and  also  because  in  roasting  and  boiling  large  pieces  of  pork  a  considerable 
time  is  necessary  to  permit  the  temperature  required  to  kill  off  the  parasites 
(62°  to  70°  C.)  to  penetrate  to  the  middle  of  the  joint,  it  appeared  to  be  most 
practical  to  have  all  pigs  microscopically  examined  for  Trichinellae  before  they,  or 
parts  of  them,  were  placed  on  the  market,  and  all  infected  meat  condemned,  no 
matter  whether  the  Trichinellae  were  present  in  large  or  small  numbers,  still 
undeveloped  or  calcified.  Since  1877  obligatory  examination  of  pork  has  been 
introduced  in  Prussia,  though  as  yet  it  is  not  thoroughly  carried  out;  other  states 
of  North  Germany  as  well  as  the  larger  towns  of  South  Germany  soon  followed  ; 
a  complete  army  of  trichina  inspectors,  officially  examined  and  periodically  con- 
trolled by  experts,  and  whose  number  in  Prussia  amounted  to  27,602  in  1896, 
this  being  even  increased  to  28,224  in  1899,  have  the  charge  of  examining 
pork  on  certain  lines  laid  down.  These  are  at  the  present  time  uniformly 
administered.  The  proceeding  is  usually  that  the  trichina  inspector  himself  goes  to 
the  slaughterhouses,  or  special  samplers  take  pieces  of  the  muscles  that  are  known 
to  be  the  favourite  seats  of  the  parasite  (pillars  of  the  diaphragm,  the  costal  part  of 
the  diaphragm,  muscles  of  the  tongue  and  larynx,  intercostal  and  abdominal  muscles) ; 
six  small  portions  are  separated  from  each  piece,  pressed  between  slides  or  special 
compressors,  and  carefully  gone  through  by  examining  them  with  a  low  power  of 
the  microscope.  The  pigs  free  from  Trichinellae  are  passed  for  commerce;  trichinous 
pigs,  on  the  other  hand,  in  Prussia,  are  only  allowed  to  be  used  for  industrial 
purposes,  i.e.^  the  hide  and  bristles  are  used,  the  fat  is  allowed  to  be  melted  down, 
or  certain  parts  are  used  for  the  manufacture  of  soap  or  glue.  In  Saxony,  however, 
it  is  still  permitted  to  place  trichinous  flesh  on  the  market,  fully  declaring  its  nature, 
and  after  having  been  heated  to  its  deepest  strata  at  a  temperature  of  100°  C.  in  a 
suitable  apparatus,  and  under  the  supervision  of  a  veterinary  surgeon. 

As  TO  THE  PROPORTION  OF  TRICHINOUS  PIGS  to  healthy  ones,  the  following 
tables  give  the  figures  for  Prussia  : — 

'  For  instance,  extensive  epidemics  occurred  in  Hettstadt  in  1863  (160  patients,  28 
deaths)  ;  Hanover,  1864 — 1865  (more  than  300  patients)  ;  Hadersleben,  1865  (337  patients, 
loi  deaths);  Potsdam,  1866  (164  patients);  Greifswald,  1866  (140  cases,  i  death); 
Magdeburg,  1866  (240  cases,  16  deaths)  ;  Halbersiadt,  1867  (100  cases,  20  deaths)  ; 
Stassfurt,  1869  (over  100  cases) ;  Wernigerode,  1873  (100  cases,  i  death) ;  Chemnitz  (194 
cases,  3  deaths);  Linden,  1874  {400  cases,  140  deaths);  Niederzwohren,  near  Cassel,  1877 
(half  the  population);  Diedenhofen,  1877  (99  cases,  10  deaths);  Leipzig,  1877  (134  cases, 
2  deaths)  ;  Ernsleben,  1883  (403  cases,  66  deaths)  ;  Strenz-Neuendorf,  1884  (86  cases,  12 
deaths),  etc.  According  to  Johne,  109  epidemics,  with  3,402  cases  and  79  deaths,  occurred 
in  Saxony  between  i860  and  1889.  Stiles,  in  a  work  recently  published,  states  that  there  were 
8,491  cases  of  trichinosis  with  513  deaths  (6*04  per  cent.)  in  Germany  from  i860  to  1880  ; 
and  6,329  cases  and  318  deaths  (5'02  per  cent.)  between  1881 — 1898.  Of  these  latter, 
1881 — 1898,  3,822  (225  deaths)  occurred  in  Prussia,  1,634  (76  deaths)  in  Saxony,  and  873 
(17  deaths)  in  the  remaining  states.  There  is,  however,  no  doubt  that  many  deaths  from 
trichinosis  were  not  recognized,  as  proved  by  experience  at  post-vwrtems. 


430 


THE   ANIMAL    PARASITES    OF    MAN 


Number  of 

Number  of 

Year 

pigs  examined 

trichinous  pigs 

Proportion 

1878 

2,524,105 

1,222 

I  :     2,065 

1879 

3,164,656 

1,938 

I  :     1,632 

1881 

3,118,780 

1,695 

I  :     1,839 

1882 

3,808,142 

1,852 

I  :    2,056 

1883 

4,248,767 

2,199 

I  :     1,932 

1884 

4,611,689 

2,624 

I  :     1,741 

1885 

4,421,208 

2,387 

I  :     1,852 

1886 

4,834,898 

2,114 

I  :    2,287 

1887 

5,486,416 

2,776 

i:     1,988 

1888 

6,051,249 

3,111 

I  •    1,945 

1889 

5,500,678 

3,026 

I  :    1,818 

1890 

5.590,510 

1,756 

I  :    3,183 

189I 

•   6,550,182 

2,187 

I  :    2,996 

1892 

6,234,559 

2,085 

I  :    2,992 

1896 

8,759,490 

1,877 

I  :    4,666 

1899 

9,230,353 

1,021 

I  :    9,040 

1902 

9,093,210 

725 

I  :  12,397 

The  proportion,  however,  is  not  only  subject  to  variation  in  separate  years,  but 
differs  according  to  the  district  ;  thus,  in  1884,  in  the  state  district  of  ?vlinden  there 
was  one  trichinous  pig  to  30,146  healthy  animals,  in  Erfurt  i  to  14,563,  in  the  district 
of  Gnesen  i  to  loi,  in  Schrimm  i  to  86,  and  in  Schroda  i  to  68. 

In  Germany  Trichinella  is  becoming  LESS  COMMON  in  pigs  (Ostertag)  :— 


{a)  Piiissia. 


Year 

1878— 1885 
1886—1892 
1896       ... 
1899       ... 
1902 


Year 

189I 
1892 

1893 
1894 

1895 
1896 

1899 
1902 


Year 
1883-1893 
1893— 1897 
1902 


{b)   Saxony. 


(c)  City  of  Berlin. 


Pigs  found 
to  be  trichinous 

o-o6i- 

-0-048  per  cent 

0-033- 

-0-043 

0-02I 

0014 

O'OII           ,, 

Num 
found  to 

ber  of  pigs 
be  trichinous 

0*014 

per  cent. 

o-oii 

,, 

o*oo8 

,, 

0-007 

,, 

0-012 

,, 

0-0I02 

,, 

0*004 
0-0056 

" 

Number  of  pigs 
found  to  be  trichinous 

0*035- 

-0*064  per  cent. 

0*022- 

-0-015       „ 

0*006 


There  is  no  doubt  that  the  excellent  preventive  measure  of 
official  inspection  for  Trichinella  has  led  to  the  avoidance  of  grave 
disasters;  its  introduction  has  not  yet  caused  an  entire  cessation 
of  trichinosis  in  man,  because  inspection  of   pork  is  not  obligatory 


DIOCTOPHYMID^  43 1 

everywhere,  so  that  human  beings  may  become  infected  by 
unexamined  trichinous  pigs  from  their  own  country  or  from  abroad^ 
and  also  because  an  infection  may  occasionally  escape  notice.  For 
these  reasons  the  meat  imported  into  Berlin  from  abroad  as  free 
from  Trichinae  is  examined  again  and  not  always  in  vain  ;  finally,  also^ 
gross  negligence  may  at  times  occur,  or  fatal  errors  may  be  made. 

In  addition  private  prophylaxis  must  not  be  neglected,  and  its 
chief  aim  should  be  directed  to  the  suitable  preparation  of  pork. 

Family.     Dioctophymldae. 
Genus.     Dioctophyme,  Collet-Megret,  1802. 

Syn.  :   Eustrongyhts,  Dies.,  1851. 

Large  worms.  Anterior  extremity  unarmed  ;  the  mouth  is  surrounded  by  six 
papillae.     One  ovary.     The  vulva  is  in  the  anterior  region   of  the  body. 

Dioctophyme  gigas,  Rudolphi,  1802. 

Syn. :  Dioctophyjjie  renale,  Goeze,  1782  ;  Ascaris  caiiis  et  mariis,  Schrank,  1788  j 
Ascaris  visceralis  et  renalis^  Gmelin,  1789  ;  Sirongylus gigas ^  Rud.,  1802  ;  Eustrongy- 
lus  <^igas.  Dies.,  1851  ;  Strongylus  renalis^  Moq.  Tand.,  i860  ;  Eustrongylus  visceralis y 
RailL,  1885. 

Colour  blood-red  ;  the  anterior  extremity  somewhat  slender ; 
there  is  a  series  of  about  150  papillae  along  the  lateral  lines ;  the 
sub-median  lines  are  strongly  developed,  and  from  them  spring  the 
radial   muscles   for  the  intestine. 

The  males  attain  a  length  of  40  cm.  and  a  diameter  of  4  to 
6  mm. ;  the  posterior  extremity  is  transversely  truncated  ;  the  anal 
orifice  is  within  the  base  of  the  collar-like  bursa,  the  thickened  edges 
of  which  are  beset  with  papillae  ;  the  spicule  measures  5  to  6  mm. 
in  length. 

The  females  attain  a  length  of  100  cm.  and  a  breadth  of  12  mm. 
The  anus  is  crescent-shaped  and  terminal.  The  vulva  is  50  to 
70  mm.  distant  from  the  anterior  extremity.  The  eggs  are  oval  and 
have  a  thick  shell  presenting  numerous  depressions  ;  the  shell  itself 
is  brownish,  but  it  is  colourless  at  the  somewhat  thickened  poles ; 
it  measures  60  //,  in  length  by  40  /a  in  breadth.  The  larva  measures 
240  /x  by  i4yL6. 

Dioctophyme  gigas  lives  in  the  pelvis  of  the  kidney,  more  rarely  in 
the  abdominal  cavity  of  the  seal,  otter,  dog,  wolf,  fox,  horse,  marten 
and  polecat,  exceptionally  also  in  human  beings.  It  also  occurs  in 
tumours  of  the  mamma  and  perinaeum.  Most  of  the  cases  in  which 
this  parasite  has  been  reported  as  occurring  in  man  may  be  traced 
back  to  unrecognized  Ascaris  hunbricoides  or  to  clots  of  fibrin ;  seven 
certain  cases,  eight  more  or  less  doubtful,  however,  remain. 


432 


THE   ANIMAL   PARASITES   OF   MAN 


The  source  of  infection  is  unknown,  but  according  to  Balbiani  the 
eggs  develop  an  embryo  in  water  or  moist  soil, 
and  this  embryo  may  remain  alive  several  years 
without  hatching ;  the  infection  of  dogs  with 
embryo-containing  eggs  did  not  succeed ;  an 
intermediate  stage  in  fishes  is  conjectured, 
but  still  the  infection  of  cattle  and  horses  is 
unintelligible. 

Famity.     Strongylidse. 
Sub-family.     Metastrongylinae,  Leiper,  1908. 

Buccal  capsule  absent  or 
slightly  developed,  vagina  elon- 
gate, uteri  convergent^  and  have 
a  simple  musculature.  Parasitic 
in  the  respiratory  or  circulatory 
system.  Genera  :  Metastrongy- 
lus,  Synthetocaulus. 

Genus.     Metastrongylus, 

Molin,  1861. 

Mouth  with  six  lips,  of 
which  the  two  lateral  are  the 
largest.  Postero-  and  postero- 
external rays-  of  bursa  thin,  the 
rest  thick.  Only  the  median 
ray  double.  Spicules  very  long 
and  slender,  striated.  Vulva 
immediately  in  front  of  anus. 
Eggs  contain  an  embryo  when 
laid. 


Fig.  306.  —  Diocto- 
phyme  gjgas,  male. 
Natural  size.  (After 
Railliet.) 


Fig.  307-— Eggs  of 
Dioctophyme  g'gas  ; 
above  seen  from  the 
flat,  below  in  optical 
section.  400/1.  (After 
Railliet.) 


Metastrongylus   apri,  Gmelin,  1789. 

Syn. :  Gordius  puhnonalis  apri,  Ebel,  I777  ;  Ascaris  apri,  Gmelin,  1789; 
Strongylus  suis,  Rud.,  1809;  Strongylus  paradoxus,  Mehlis,  1831  ;  Strongylus 
elongatus,  Duj.,  1845;    Strongylus  longevaginatus.  Dies.,   185 1. 

The  male  measures  12  to  25  mm.  in  length  ;  the  bursa  is  bilobed  ; 
there  are  five  rays  in  each  lobe ;  the  spicules  are  thin  and  up  to 
4  mm.  in  length.  The  females  measure  20  to  50  mm.  in  length, 
the    anus    is    close    in    front    of    the     posterior    extremity,    which 

*  Convergent :  i.e.,  the  uteri  are  parallel,  converging  from  the  anterior  part  of  body  to  the 
vagina,  which  is  near  the  anus,  this  position  being  associated  with  convergence  of  the  uteri. 
Divergent :  Uteri  run  anterior  and  posterior,  diverging  from  the  vagina,  which  in  this  case  is 
near  middle  of  body, 

''■  For  nomenclature  of  rays  vide  p.  449. 


TRICHOSTRONGYLIN^ 


433 


CL.yrt 


has  a  recurved,  hook-like  process  ;  the  vulva  is  close  in  front  of  the 
anus.  The  eggs  are  elliptical,  57//,  to  100  yit  in  length,  39  /a  to  72  yu,  in 
breadth  ;  when  the  eggs  are  deposited  the  embryo  is  already  formed, 

220//,  to  350//,  by  lOyLt  to   12  fJb. 

Metastrongylus  apri  frequently  lives  in  the  bronchial  tubes — usually 
the  smaller  ones — of  the  pig^  and  wild  boar  ;  it  is  also  found  occa- 
sionally in  sheep  and  in  man  ;  in  young  pigs  it  is  apt  to  set  up 
a  bronchitis,  which   frequently  causes  death. 

The  first  communication  as  to  the  occurrence  of  this  species  in 
man  was  that  of  Diesing,  who,  in  1845,  in  Klausenburg,  had  the 
opportunity  of  examining  Strongylidce  found  by  Jortsits  in  the  lung 
of  a  little  boy,  aged  6,  in  Transylvania;  probably  also  the  Nema- 
todes found  in  the  trachea  and 
larynx  of  man,  and  described  by 
Rainey  and  Bristowe  as  specimens 
of  Filaria  tracliealis^  belong  to 
this  group ;  according  to  Chatin, 
Metastrongylus  apri  may  also  occur 
in  the  intestine  of  man ;  this  oc- 
currence, however,  may  in  all 
probability  have  been  due  to  an 
accidental  introduction  of  adult 
worms  into  the  intestine,  and 
should  not  be  attributed  to  an 
infection  by  the  larval  stage. 

No  experiments  to  induce  in- 
fection have  been  made ;  it  is  prob- 
able, however,  that  infection  is 
direct  and  without  the  aid  of  an 
intermediate  host. 


Fig.  308. — Metastrongylus  apri:  one  side 
of  bursa,  a.,  anterior;  a.e.^  antero-external ; 
a.m.,  antero-median  ;  p.m.,  postero-median  ; 
p.e.,  postero- external  ;  /. ,  one  division  of 
posterior  ray.     (Stephens.) 


Sub-family.     Trichostrongylinae,  Leiper,   1908. 

Strongylidae  with  buccal  capsule  absent  or  slightly  developed, 
vagina  short,  uteri  divergent  (/.^.,  anterior  and  posterior),  ovejectors 
differentiated.  Parasitic  in  the  alimentary  canal.  Contains  the 
genera  Trichostrongylus,  Haemonchus,  Ostertagia,  Nematodirus, 
Cooperia,  Dictyocaulus.'^ 

1  The  reports  of  the  city  inspection  of  meat  in  Berlin  state  that  Strongylidce  in  the  lungs 
of  pigs  are  by  no  means  rare;  therefore  the  lungs  of  1,941  pigs  were  condemned  between 
1885— 1886,  of  1,641  between  1886— 1887,  of  3,237  between  1887—1888,  of  4,855  between 
1888 — 1889,  of  7,197  between  1889— 1890,  and  of  5,574  pigs  between  1890— 1891,  etc. 
Ostertag  found  Stro7igyliis  apri  in  60  per  cent,  of  the  pigs  examined  in  the  Berlin  abattoir  ; 
Meyer,  in  Leipzig,  found  the  parasite  in  15  per  cent,  of  the  native  pigs  and  in  52  per  cent,  of 
the  Hungarian  pigs. 

^  Dictyo:aidns  is  parasitic  in  the  bronchi.  ' 


434 


THE   ANIMAL   PARASITES   OF   MAN 


Genus.     Trichostrongylus,  Looss,  1905. 

Very  small  Strongylidce,  Mouth  with  three  small  lips  and  nodular 
or  punctiform  papillae.  Cervical  papillae  absent.  Bursa  entirely 
closed;  with  large  lateral  lobes,  and  median  lobe  not  distinctly  defined. 
Anterior^  rays  double,  the  branches  widely  divergent,  one  thin, 
the  other  thick,  and  close  to  the  antero-median.  The  postero- 
median ray  is  thin  and  close  to  the  postero-external.  Posterior  ray 
bifurcate,  each  branch  bifid  at  the  tip  (fig.  311).  Spicules  short,  spoon 
or  spatula-like,  with  on  the  broad  anterior  end  a  lateral  knob  or  disc 
and  in  h'ont  of  the  point  an  angular  projection.  Gubernaculum  of  a 
peculiar  canoe  or  shoe  shape  in  profile.  Vulva  in  the  hinder  half  of 
the  body.  Tail  with  two  minute  papillae  just  in  front  of  tip.  Eggs 
thin  shelled ;  when  laid  they  show  eight  to  thirty-two  segments. 
Parasitic  in  duodenum,  seldom  in  the  stomach  of  herbivora. 


Syn. 


Trichostrongylus  instabllis,^  Railliet,  1893. 
Stro7igylus  instabilis^  Railliet,  1893;  Strongylus  subtilis,  Looss,  1895. 


Male  4  to  5*5  mm.  long,  o'o8  mm.  thick  in  front  of  bursa. 
Spicule  0*135  to  o'i45  mm.  long,  accessory  piece  (gubernaculum) 
0*07  mm.  thick.  Antero-external  ray  usually  thickest  of  all,  occa- 
sionally only  as  thick  as  the 
antero-median ;  postero-median 
far  more  slender  than  the  antero- 
external  and  antero-median  and 


; 


Fig.  309. — Trichostrongylus  instabilis  :  left, 
posterior  end  of  male  ;  right,  spicule  and 
gubernaculum,  side  view.  C/.  fig.  311.  Mag- 
nified.    (After  Looss.) 


Fig.  310. — Trichostrongylus  instabilis  : 
posterior  end  of  female.  Magnified. 
(After  Looss.) 


nearer  to  the  postero-external  than   to  the   antero-median.     Female 
5  to  6  mm.  long,  vulva  1-05  to  1*2  mm.  distant  from  the  tip  of  the 

•  When  the  anterior  ray  is  double,  the  branches  of  it  are  called  an tero -anterior  and  latero- 
anterior. 

^  Identical  with  T.  colubriformis  of  the  sheep  according  to  Leiper.  If  so,  this  latter  name 
has  priority. 


TRICHOSTRONGYLUS    PROBOLURUS 


435 


tail,  placed  longitudinally,  50  /x  to  55  /a  long,  always  shorter  than 
the  unpaired  portion  of  the  canal  formed  by  the  union  of  the  two 
ovejectors  ;  anus  0*055  to  0*07  mm.  distant  from  tip  of  the  tail ; 
ova  73  yLt  to  80  /A  by  40  /a  to  43  yu,. 

This  species  lives  in  the  duodenum,  exceptionally  also  in  the 
stomach  of  Oris  aries,  0.  laficaudn,  Ant  Hope  dorcas,  Camelus  dronie- 
darius  (Egypt),  CynocepJialns  haniadryas  (North  Africa),  sheep  and 
goats  (France),  and  has  been  found  by  Looss  in  bodies  of  fellaheen 
at  Alexandria  and  in  the  stomach  of  a  Japanese  female  by  Ijima. 


Trichostrongylus  probolurus,  Railliet,  1896. 

Syn.  :  Strongylus  probolurus^  Railliet,  1896. 
Male  45  to  5'5  mm.  long,  in  front  of  bursa  o'o8  mm.  thick ; 
spicule  o'i26  to  o'i34  mm.  long,  gubernaculum  0*075  to  o'o8  mm, 
long.  Bursa  :  latero-anterior  rib  thickest ;  antero-external  thicker 
than  antero-median,  postero-median  and  postero-external  very  short 
and    close   together.      Female   4*5    to    6   mm.  long,    vulval    opening 

I'oS  to  1*25  mm.  from  tip 
of  tail,  placed  longitudinally , 
and  slightly  curved,  76  /a  long. 


Fig.  312. —  Trichostrongylus  probolurus ." 
spicules  and  gubernaculum  of  male  ;  on 
left,  ventral  view  ;  on  right,  lateral  view. 
X  c.  300.     (After  Looss.) 


Fig.  311.  —  Trichostrongylus  probolurits  :  tail  of 
male  from  left  side,  d.,  posterior;  e.d.,  postero- 
external ;  /./.,  postero-median  ;  m.l.,  antero- 
median ;  e.i.,  antero-external;  /.z/.,  latero-anterior; 
V.V.,  antero-anterior  ;  g-zib.,  portion  of  guberna- 
culum ;  sp.,  portion  of  spicules.  x  c.  300. 
(After  Looss.) 

always  longer  than  the  unpaired  portion  of  the  ovejector  ;  anus  0*040 
to  0*05  mm.  distant  from  tip  of  tail.  Posterior  end  thick,  point  of  tail 
short.     Ova  76  //,  to  80  yu,  by  43  yu,  to  46  /ju. 

Habitat. — In  the  duodenum  of  Ovis  aries,  0.  laticatida,  Antilope 
dorcas,  Camelus  droniedarius  (Egypt)  and  occasionally  also  in  man 
(Egypt). 

Trichostrongylus  vitrinus,  Looss,  1905. 

Male  4  to  5*5  mm.  long,  in  front  of  bursa  0*085  mm.  thick. 
Bursa  larger  than  in  the  other  two  species,  antero-external  rib  thickest, 

28 


436 


THE   ANIMAL  PARASITES   OF  MAN 


antero-anterior  and  postero-median  equally  thick,  straight.  Spicule 
o*i6  to  0*17  mm.  long,  gubernaculum  0*085  to  0*095  ^^'  long- 
Female  5  to  6-5  mm.  long,  vulval  opening  1*15  to  1*25  mm.  distant 
from  tip  of  tail,  crescent  shaped,  oblique  to  body  axis,  and  around 
it  irregular  thickenings.     Ova  84  yit  to  90  /x  by  46  fi  to  50  //,. 

In  duodenum   of  Ovis 

aries,    0.  laticatida,  occa- 

(7u^if.      sionally  in  Camelus  dronie- 

dariiis  and  in  man  (Egypt). 


v.v- 


^-&.d. 


YiG.  ZiX.—  Trickostrofjgylus  vitrinus  :     tail  of  male  Y\G.Z\i^.-Trichostrongylus 

from  left  side.      «'.,   posterior  ;    e.d.,    postero-external ;  z'^/rz^z.^  :  spicules  and  guber- 

/./.,  postero-median  ;  ^^/./.,  antero-median;  ^./.,antero.  naculum ;    on     left,     ventral 

external;    l.v.,    latero-anterior  ;    v. v.,   antero-anterior;  ^^^w  ;  on  right,  lateral  view. 

gub.,  portion  of  gubernaculum  ;  sp.^  portion  of  spicule.  ^  ^'  3°°'     (Alter  Looss.) 
X  c.  300.     (After  Looss.) 

Genus.     Haennonchus,  Cobb.,  1898. 

Small  mouth  cavity  contains  a  ''  tooth  "  or  ^*  lancet "  arising  from 
the  dorsal  side.  Cuticle  of  head  and  neck  not  inflated.  Cervical 
papillae  well  marked.  Bursa  bilateral,  with  large  lateral  lobes  and  a 
small  dorsal  lobe  not  median,  but  lateral,  attached  to  the  base  of  one 
of  the  lateral  lobes  (fig.  316).  Posterior  ray  bifurcate,  each  branch 
tifid  apically.  Each  lateral  lobe  six  rays.  Anterior  rays  separated 
distally,  curving  forward.  Antero-median  and  postero-median  rays 
distally  curve  away  from  the  antero-external.  Postero-external  ray 
long  and  slender.  Spicules  less  than  i  mm.  Gubernaculum  present. 
Vulva  in  posterior  part  of  body  covered  by  a  prominent  tongue-like 
flap.     Eggs  ellipsoidal. 

Haemonchus   contortus,  Rudolphi,  1803  ;   Cobb.,  1898. 

Dorsal  ''tooth"  or  "lancet"  10 /x  to  i5yLt  long.  Cervical  papilla 
C3  mm.  from  head. 

Male  20  mm.  long  by  400  /x  thick  (maximum).    Asymmetrical  lobe 


H^MONCHUS   CONTORTUS 


437 


of  bursa  150  yu,  by  125  //,  attached  to  left  lateral  lobe.  Posterior 
ray  bifurcate  ;  each  branch  bifid.  Stem  of  ray  less  than  twice  as  long  as 
its  branches.  Spicules  300  yu,  to  500  /^  with  knobbed  tips,  and  the  left 
spicule  with  a  barb  20  fju  from  the  tip,  right  spicule  with  a  barb  40  /a 

from  tip.  Gubernaculum  200  />«,  by 
25  yL6  to  35  /x,  fusiform  with  thick- 
ened edges. 

Female  18  to  30  mm.  by  500  fj. 
(maximum).  Vulva  3  to  4*5  mm. 
from  tip.  Linguiform  flap  0*5  mm. 
(a  second  one  exists,  according  to 
Brumpt).  Anus  400  fi  to  630  //,  from 
tip.   Tail  acutely  pointed.    Eggs  75  /a 

to  95  yU<  by  40  yLt  to   50  fJL. 


:^;..'OV^ 


— int. 


A — e^eL 


Fig.  315.  —  Hcemonchus  contortus: 
vulval  region  of  female  viewed  from 
left  side,  int.^  intestine;  lab.^  lingui- 
form process  covering  vulva  ;  ov.^  ovary  ; 
ovij.y  ovejector ;  ut.^  uterus;  vag., 
vagina ;  z/«/.,  vulva.  X  75.  (After 
Ransom.) 


m.l. 


Fig.  316.  —  Hcemonchus  contortus:  tail  of 
male,  dorsal  view,  d.,  posterior  ray  of  the  asym- 
metrically placed  posterior  lobe;  ^.o'.,  postero- 
external ;  /./.,  postero-median ;  mj.,  antero- 
median ;  <?./.,  antero  -  external  ;  l.v.,  latero- 
anterior ;  v.v.,  antero-anterior ;  gub.,  guberna- 
culum; ^/.,  spicule.     X  75.     (After  Ransom.) 


Habitat. — Fourth  stomach  of  cattle,  sheep,  antelope. 
Distribution.  —  Europe,    America,    Africa,    Asia,    Australia,    New 
Zealand.     Once  in  man  in   South  America  by  de  Magalhaes. 

Pathology. — Produces  anaemia,  emaciation,  dropsy  in  sheep  ;  and 


438  THE   ANIMAL   PARASITES   OF   MAN 

in    the    human    case    the    symptoms   were    mistaken    for   those   of 
ancylostomiasis. 

Life-history.  —  Rhabditic  embryos  easily  hatch  in  water,  then 
moult  several  times,  becoming  eventually  *^  filariform  "  larvae  enclosed 
in  the  moulted  skin.  These  crawl  up  blades  of  grass  and  are 
swallowed  by  sheep,  etc. 

Genus.     Nematodirus,  Ransom,  1907,  emend.  Railliet,  191 2. 

Head  over  50//,  in  diameter.  Cuticle  may  be  slightly  inflated  and 
often  transversely  striated.  Cuticle  with  eighteen  distinct  longi- 
tudinal ridges.  Cervical  papillae  absent  (?).  Posterior  lobe  of  bursa 
reduced  to  short  lobules  each  with  a  dorsal  ray.  Antero-anterior 
4-  latero-anterior  (=  anterior  double)  rays  close  together,  parallel ; 
antero-external  ray  diverges  widely  from  antero-  and  postero-median, 
which  are  close  together  and  parallel.  Postero-external  ray  slender. 
Spicules  more  than  0*5  mm.  long,  at  most  one-twelfth  of  body,  united 
by  a  membrane'  throughout  their  length  or  only  distally.  Guber- 
naculum  absent.  Vulva  behind  middle  of  body.  Eggs  ellipsoidal, 
shell  rather  thick. 

Habitat. — Duodenum  of  ruminants. 

Sub-genus.     Mecistocirrus,  Railliet,  191 2. 

Head  slightly  inflated,  with  transverse  striations.  Skin  with 
eighteen  longitudinal  ridges,  but  little  apparent ;  cervical  papillae 
distinct.  Bursa  bilobed  ;  median  ray  double  (=  postero-median  + 
antero-median) ;  very  large  antero-external  at  the  edge,  close  to  the 
anterior.  Spicules  very  long,  slender,  one-sixth  length  of  body 
(3-5  mm.) ;  tail  pointed.     Vulva  immediately  in  front  of  anus. 

Habitat. — Stomach  of  ruminants. 

Mecistocirrus  fordi,  Daniels,  1908. 

Syn.  :  Strongylus fordi,  Daniels,  1908;  Strongylus  gibsoni^  Stephens,  1909; 
Nematodirus  fordi,  Leiper,  191 1. 

Male  21  mm.  long  by  0-4  mm.  thick.  Cervical  papillae  045  mm. 
behind  the  head.  Spicules  about  7  mm.  long,  i.e.,  one-third  of  the 
body  length.  At  the  level  of  the  postero-external  rays  of  the  bursa, 
the  bursa  has  a  projecting  lobule. 

Female  25  mm.  long.  Anus  0*2  mm.,  vulva  0-5  mm.  from  the  tip 
of  tail.     Eggs  100 /x  by  53 /x. 

Sub-family.     Ancylostominae,  Railliet,  1909. 
Strongylidce  with  buccal  capsule,  well  developed.     Uteri  divergent. 
Parasitic  in  the  alimentary   canal,   exceptionally   in    the   respiratory 
system. 


GESOPHAGOSTOME^ 


439 


Group.    (Esophagcstomese,   Railliet  and  Henry,  1909. 

Bursa  with  anterior  and  median  ray  cleft  (not  double),  postero- 
and  postero-external  arising  from  a  common  trunk,  posterior  bifur- 
cated, each  limb  bi digitate. 

Contains  at  present  four  genera  :  (i)  Ternidens,  (2)  Chabertia, 
^3)  Qi^sophagostomum,  (4)  Agriostomum. 


Fig.  317. — Mecistocirrus  forat .  bursa  of  male,  dorsal  view.  The  rays 
are  (i)  postero-external,  (2)  median  (=  postero-median  +  antero-median),  (3) 
antero-external,  (4)  latero-anterior,  (5)  antero-anterior.  These  two  latter  are 
parallel.     The  posterior  ray  is  absent.     (After  Stephens.) 

Genus.     Ternldens,  Railliet,  1909. 

Buccal  capsule  sub-globular,  opening  obliquely  in  the  dorsal  surface, 
and  having  at  the  bottom  three  complex  teeth  resembling  those  of 
Triodontophorus.^  Two  crowns  of  leaflets  ;  peristomic  collar  moderate, 
edge  of  bursa  slightly  toothed. 

Type. — r.  deminiituSy  Railliet  and  Henry, 

1  Triodontophorus  belongs  to  the  group  CylicostomecE,  which  has  the  following  bursal 
formula:  (i)  anterior  cleft,  (2)  median  double,  (3)  postero-external  and  posterior  arising 
separately,  (4)  posterior  double,  each  branch  giving  off  two  lateral  branches. 


440 


THE   ANIMAL   PARASITES   OF   MAN 


Ternidens  deminutus,  Raill.  and  Henry,  1905. 
Syn.  :   Triodontophorus  deminutus,  Raill.  and  Henry,  1905. 
Body  relatively  thick.     Cervical  papillae  0*5  mm.  behind  the  head. 
Buccal  capsule  40  y.  deep.     Teeth  40  /i  long. 


Fig.  318.  —  Ternidens  detnimitus.  A,  head  end,  ventral  view:  c,  crown  of 
leaflets;  v.o.,  buccal  cavity;  d,  pharyngeal  plates;  ph.,  pharynx;  n.,  valve. 
B,  lateral  view.  C,  tail  of  female.  D,  bursa  of  male  :  a.,  anterior  ray  ;  a.e., 
antero-external  ;  m.,  median;  p.e.,  postero-external ;  p.,  posterior.  E,  pharyngeal 
plate.     Enlarged.     (After  Railliet  and  Henry.) 

Male  9-5  mm.  long  by  560  fju  thick.  CEsophagus  660  /jl  long. 
Bursa  broader  than  long,  the  lateral  lobes  united  by  a  small  posterior 
lobe  with  slightly  sinuous  margin  ;  edge  of  bursa  finely  toothed. 
Spicules  about  900  /ul  long. 


(ESOPHAGOSTOMUM  44I 

Female  12  to  16  mm.  long  by  650  fju  to  730  fx  thick.  (Esophagus 
860  fjL  long.  Vulva  forms  a  distinct  projection  480  //,  from  tip  of  tail. 
Anus  240  yLt  to  270/1,  from  tip.     Eggs  60  //,  to  80  //,  by  40  /a. 

Habitat. — Large  intestine  of  a  negro  (Comoro  Islands)  and  in  the 
natives  of  Nyasaland  and  Portuguese  East  Africa.  Also  in  large 
intestine  of  Macacus  sinensis  and  Macaciis  cynoinolgus. 

Genus.     CEsophagostomum,  Molin,  1861. 

No  teeth.  Cuticle  around  the  mouth  dilated  to  form  a  narrow 
cuticular  "  peristomic  collar."  Separated  by  a  constriction  from  this 
is  a  much  more  extensive  inflation,  the  "  cephalic  vesicle,"  bounded 
abruptly  behind  on  the  ventral  side  by  a  transverse  groove,  the 
"  ventral  cleft,"  which  is  always  present  even  in  absence  of  the  vesicle. 
Buccal  cavity  of  slight  depth  with  a  short  dorsal,  tunnel.  Internal 
margin  of  the  mouth  armed  with  chitinous  leaflets  ("external  crown"); 
internal  border  of  the  buccal  capsule  armed  with  short  tongue-like 
leaflets  (internal  crown).  Lateral  membranous  wings  may  extend 
backwards  from  the  ventral  cleft.  Cervical  papillae  present.  Bursa 
with  two  lateral  lobes  united  by  a  smaller  median  lobe.  Spicules 
over  5  mm.  long,  slender ;  gubernaculum  inconspicuous.  Vulva 
in  front  of  anus.  Adults  usually  in  large  intestine  of  ruminants, 
suidae,  tapirs,  edentates  and  apes.  Larvae  sometimes  in  nodules  in 
intestinal  wall. 


CEsophagostomum   brumpti,  Railliet  and  Henry,  1905. 

Female  immature,  8*5  to  10*2  mm.  long,  0*295  to  0*325  mm. 
thick.  Cuticle  transversely  striated.  The  cephalic  vesicle  imme- 
diately behind  the  vestibulum  oris,  embracing  the  anterior  two-fifths 
of  the  oesophagus,  extending  ventrally,  however,  towards  its  posterior 
end.  Vestibulum  oris  formed  by  a  cuticular  band  provided  with  a 
crown  of  twelve  apical  leaflets  directed  forward  and  inwards  ;  six 
cephalic  papillae  (two  lateral,  four  submedian) ;  buccal  capsule  in  front 
of  cervical  swelling  not  delineated  circularly  behind,  but  provided 
with  three  wide  incisions  (one  dorsal,  two  sub-ventral).  (Esophagus, 
0*470  to  0*500  mm.  long,  two  cervical  papillae  at  five-eighths  of 
its  length.  Vulva  0*350  to  0*475  mm.,  anus  0*170  to  0*200  mm.^ 
before  tip  of  tail. 

Habitat. — Found  by  Brumpt  in  tumours  of  the  caecum  and  colon 
of  a  native  of  the  River  Omo  (Lake  Rudolph),  East  Africa.  Immature 
forms  only  were  present.  Adults  have  been  found  in  similar  tumours 
in  monkeys. 

Pathology. — They  occur  in  haemorrhagic  cysts  in  the  submucosa  or 


442 


THE   ANIMAL   PARASITES   OF   MAN 


.AM>o^u:d^ 


Fig.  319. —  (Esophagostornum  stephatiostomum  var.  thomasi.  i,  male,  natural  size  ;  2, 
female,  natural  size;  3,  head  of  female,  ventral  view,  showing  cephalic  vesicle  and  ventral  cleft 
limiting  it  behind,  x  55  ;  4,  head  of  female,  dorsal  view,  x  225  ;  5,  head  of  male,  end  view, 
showing  external  and  internal  leaf  crowns,  x  225  ;  6,  tail  of  male,  lateral  view  [cf.  fig.  318,  D), 
X  20;  7,  tail  of  female,  lateral  view,  x  20  ;  8,  CEs.  thomasi,  posterior  ray  of  bursa,  x  150; 
9,  (Es.dentigerum,  from  chimpanzee,  posterior  ray  of  bursa,  X  150;  10,  CEs.  stephanostomum, 
from  gorilla,  posterior  ray  of  bursa,   x   150. 


GESOPHAGOSTOMUM   STEPHANOSTOMUM 


443 


muscularis  mucosae  of  the  gut  wall.  The  cysts  project  internally  and 
externally,  and  contain  inmature  adults,  which  eventually  escape  into 
the  lumen  of  the  gut. 


Fig.  320. — CEsophagosiomum  stephanostomum  var.  thomasi :  csecum  and  ascending  colon. 
Subperitoneal  cysts  are  seen  on  the  top  right  hand,  and  in  the  lumen  of  the  gut  numerous 
cysts  arranged  transversely.  The  small  roundish  patches  are  areas  of  necrosis  in  the  cyst  walls. 
^ After  Thomas.) 

CEsophagostomum  stephanostomum  var.  thomasi, 
Raill.  and  Henry,  1909. 

Body  thick,  pointed  only  at  the  ends.  Buccal  capsule  much 
reduced.  External  crown  of  thirty-eight  leaflets  (the  *^  crown  "  nearest 
the  centre  of  lig.  319,  5).  Male  17  to  22  mm.  long  by  750  //,  thick. 
Spicule    1*380   to    1*475    mm.,    slightly    curved   at   the   tip.     Female, 


444 


THE   ANIMAL   PARASITES   OF   MAN 


iinmature,  i6  to  20  mm.  long  by  900  yu,  thick,  tail  ending  in  a  little 
conical  appendage.  Anus  230  jju,  vulva  500  jx  to  525  fi  from  tip. 
Ovejectors  close  together.     Uteri  very  short  in  form  of  oblong  pouch, 

CEs.  stephanostomuni,  Stossich,  1904,  m  the  large  intestine  of  gorilla. 
(Es.  stepliauostomuui  var.  dentigera,  Raill.  and  Henry,  1909,  in  the 
chimpanzee. 

Habitat. — In  large  and  small  intestine  of  man,  Brazil. 

Pathology. — Nodules  occur  in  the  gut  wall ;  187  were  found  by 
Thomas  in  his,  the  sole  case.    The  tumours  contain  each  a  single  worm. 


Fig.  321.  —  (Esophagoslomuvi  Stephanos totmim 
var.  thomasi :  portion  of  the  ileum,  showing  a  cyst 
with  protruding  worm,      x  8.     (After  Thomas.) 


Fig.  322.  —  CEsophagostomum  ste- 
phanosto77iuvi  var.  thomasi  :  colon  with 
oesophagostome  withdrawn  from  its  cyst 
cavity,      x  20.     (After  Thomas.) 


CEsophagostomum  apiostomum,  Willach,  1891. 

According  to  Leiper,  CEs.  hnimpti  is  identical  with,  and  hence  a 
synonym  of,  this  species.  Parasitic  in  large  intestines  of  monkeys, 
producing  dysentery,  and  in  man  (Northern  Nigeria). 

According  to  Walker  this  species  is  common  in  Philippine  monkeys. 
Ova  are  scanty  in  the  faeces.  They  measure  73  /a  to  84  //,  by  44  /x  to 
57  fi  and  are  in  the  morula  stage.  They  are  easily  cultivated.  The 
rhabditiform  larva  is  340  yLt  by  16  /x  and  has  a  long  filiform  tail.  It 
moults  twice,  and  at  the  second  moult  becomes  a  filariform  larva 
retaining  the  skin  of  this  moult,  this  stage  being  that  of  the  mature 
larva.  It.now  measures  9  mm.  long  by  30  /x  thick.  Walker  suggests 
that  the  mode  of  infection  is  similar  to  that  of  ancylostomes. 


ANCYLOSTOME^  445 

Group.    Ancylostomeae,  Railliet  and  Henry,  1909. 

Bursa  with  anterior  ray  cleft,  median  double,^  postero-  and  postero- 
external arising  from  a  common  trunk,  posterior  bifurcate,  each  limb 
being  tridigitate.     Vulva  in  posterior  third  of  body.     Uteri  divergent. 

Contains  the  following  genera  :  (i)  Strongylus,'  (2)  Ancylostoma, 
(3)  Uncinaria,  (4)  Characostomum,  etc. 

Genus.     Ancylostoma,  Dubini,  1843,  emend.  Looss,  1905. 

Ventral  margin  of  mouth  capsule  armed  with  teeth,  the  "roots" 
of  which  are  continued  backwards  and  appear  on  the  external  surface 
of  capsule  as  rib-like  thickenings.  Terminal  third  of  dorsal  ray 
cleft.  Genital  tubes  very  long,  with  short,  closely  packed  diagonal 
convolutions. 

Ancylostoma   duodenale,  Dubini,  1843. 

Male  9  mm.  long  by  0*45  mm.  thick,  female  12  mm.  long  by 
0*6  mm.  thick.  Pale  flesh  colour,  or  an  intense  red  in  posterior  third. 
Anteriorly  may  be  more  or  less  black  due  to  (blood)  pigment  in  the 
cells  of  the  chyle  intestine  (=  stomach  +  small  intestine).  The  worm 
is  about  the  same  thickness  all  through  and  is  plump  and  rigid. 
Cuticle  striated.  The  body  has  a  marked  torsion,  so  that  if  the  ventral 
side  of  the  head  is  upwards  the  anus  appears  to  open  laterally  and 
vice  versa.  The  dorsal  curve  of  the  head  end  is  only  slight  and 
the  oesophagus  is  roughly  cylindrical. 

Buccal  Capsule. — The  buccal  capsule  is  bent  dorsally,  0*2 1  mm. 
long,  0'i9  mm.  broad.  If  a  worm  is  rolled  under  the  cover-glass  so 
that  the  dorsal  side  is  upwards,  we  observe  the  following  features 
(fig.  325)  :  In  the  dorsal  edge  of  the  chitinous  capsule  there  is  a  gap 
as  if  a  U-piece  had  been  punched  out.  This  is  the  "  dorsal  gap  or 
incision."  The  so-called  ^'  dorsal  teeth "  are  simply  the  rounded 
edges  of  the  tips  of  this  gap.  They  project  beyond  the  skin  which 
covers  the  capsule  externally.  Below  this  gap  is  seen  a  curved  line 
which,  if  followed  along  the  sides  of  the  capsule  on  each  side,  merges 
into  the  base  of  the  most  posterior  ventral  tooth.  This  line  is  the 
optical  expression  of  a  very  shallow  groove  on  the  inside  of  the 
capsule.  The  skin  on  the  outside  of  the  capsule,  which  is  reflected 
over  the  edge  of  and  into  the  capsule,  dips  into  this  groove,  which 
gives  it  a  firm  attachment.  Below  the  middle  (dorsally)  of  this  curved 
line  there  is  a  thickening  in  the  capsule  wall,  which  is  perforated  by  the 
opening  of  the  dorsal  oesophageal  gland.  This  is  the  ''dorsal  ridge" ;  in 
optical  section  it  has  a  conical  appearance  with  a  lumen  (of  the  duct). 

'  I.e.^  with  a  distinct  space  between  the  limbs. 

*  Strongylus  (Syn.  :  Sclerostomum)  differs  slightly  in  its  posterior  ray  from  the  other 
genera  of  the  group.     Each  bifurcation  is  trifurcate  rather  than  tridigitate. 


446 


THE   ANIMAL   PARASITES   OF   MAN 


On  the  ventral  wall  one  sees  the  two  pairs  of  strong  teeth,  their 
points  being  directed  somewhat  backwards.  They  are  covered  by 
cuticle  above  and  below,  but  their  points  are  free,  piercing  the  cuticle. 


Gh 


Cdr 


Sp 


Gcph 


Ut 


I. 


^ 


Bni- 


Fig.  323.  —  Ancylostonia  duodtjiaie, 
male.  B,  bursa ;  Bm,  bursal  muscles ; 
Cdr^  cement  gland  surrounding  the  ejacu- 
latory  duct  ;  Glc,  cervical  glands ;  N, 
nucleus  of  cephalic  gland  ;  AV,  nerve  ring  ; 
Ty  testes ;  Sp,  spicule  ;  Vs,  vesicula  semina- 
lis.     X  15.     (After  Looss.) 


Fig.  324.  —  Ancylostoma  duodenale, 
female.  Ay  anus  ;  Gcph,  cephalic  gland  ; 
Ny  nucleus  of  cephalic  gland  ;  Glc,  cervical 
gland  ;  Ov,  ovary  ;  Pex,  excretory  pore ; 
Rs,  receptaculura  seminis  ;  Ui,  uterus  ; 
F,  vagina,      x   15.     (After  Looss.) 


The  "roots"  of  these  teeth  followed  backwards  appear  as  two 
thickenings  or  ribs  on  the  outside  of  the  capsule  w^all,  so  that  the 
outside  wall  is  not  smooth — a  characteristic  of  the  genus  Ancylo- 
stoma.    In  the  space  between  these  ribs  lies  the  ventral  nerve  papilla, 


ANCYLOSTOMA    DUODENALE 


447 


and  lying  against  the  outside  of  the  outer  root  the  lateral  nerve 
papilla.  The  nerve  papillae  are  thus,  as  it  were,  concealed  by  these 
roots,  and  not  conspicuous  as  they  are  in  Necator.  Following  the 
ventral  curve  of  the  capsule  on  the  inside,  posteriorly  we  next  find 
two  triangular  ventral  lancets.^  These  stand  straight  up  into  the 
capsule  on  either  side  of  the  longitudinal  axis,  converging  at  their 
summits.  So  that  to  sum  up,  the  cutting  apparatus  is  entirely 
ventral,  consisting  of  two  pairs  of  cutting  teeth  and  a  pair  of  lancets. 

Cervical  Papillcv. — Two,  one  on  each  side  behind  the  head  at  the 
level  of  the  excretory  pore.     They  consist  of  ''pulp,"  i.e.,  extensions 
of  the  substance  of  the  lateral 
bands  covered  by  cuticle   and 
supplied  with  a  nerve  (fig.  326). 

CEsopJiageal  Glands  (3).  — 
The  chitin  of  the  triradiate 
oesophagus  is  continuous  with 
that  of  the  buccal  capsule.  In 
its  muscular  walls  are  three 
glands  —  one  dorsal,  two  sub- 
ventral.  The  dorsal  gland  opens 
into  the  buccal  cavity  through 
the  dorsal  ridge ;  the  two  others 
into  the  lumen  of  the  oeso- 
phagus at  the  nerve  ring.  They 
branch  freely  amidst  the  muscle? 
They  are  probably  digestive  in 
function. 

Cephalic  Glands  (2). — Lie  in 
the  lateral  lines  or  bands  on 
either  side.     They  begin  about 

the  middle  line  of  the  body,  and  their  ducts  open  at  the  base  of  the  outer 
ventral  tooth  on  the  surface  of  the  skin  on  each  side.  Each  is  0*15  mm. 
thick  in  the  middle,  and  has  a  single  nucleus  about  as  big  as  an  ancy- 
lostome  egg  (N,  fig.  323).     They  probably  function  as  poison  glands. 

Excretory  System  and  Cervical  Glands  (2). — The  excretory  pore  lies 
in  the  mid  line  ventrally  behind  the  oesophageal  nerve  ring  (figs. 
324  and  326).  It  opens  into  the  excretory  vesicle,  a  cavity  in  a  large 
cell  with  lateral  appendages  which  fuse  with  the  lateral  lines,  this 
cell  thus  forming  the  ''  bridge  "  of  the  excretory  system.  Adhering 
to  this  (bridge)  cell  are  the  spindle-shaped  cervical  glands  (G/c, 
fig.  324),  and  branches  from  the  excretory  vesicle  enter  the  glands, 
which  are  excretory  in  function  ;  the  vesicle  also  receives  branches 


Fig.  325. — Ancylosto7na  duodenal e  :  showing 
ventral  teeih,  dorsal  cleft,  and  behind  it  the  dor- 
sal ridge  with  duct  of  dorsal  oesophageal  gland. 
X  c.  200.     (After  Looss.) 


The  ventrallancet  (of  one  side)  of  Necator  is  seen  in  fig.  335. 


448 


THE   ANIMAL   PARASITES   OF   MAN 


from  the  lateral  excretory  canals  (fig.  326)  running  in  the  lateral 
lines  or  bands.  The  cervical  glands  are  swollen  anteriorly,  forming 
the  so-called  ampullae  just  in  front  of  the  bridge.  They  extend 
backwards  a  little  beyond  the  anterior  loop  of  the  testis. 

Lateral  Lines. —  (i)  Are  broad  elevations  of  the  subcuticle,  in 
which,  here  and  there,  a  nucleus  occurs.  (2)  Near  the  bursa  in  the 
male  they  increase  in  volume,  and  finally  divide  into  branches  which 
form  the  "pulp"  of  the  different  rays.  (3)  In  addition  to  the  lateral 
lines  or  bands,  there  is  also  a  dorsal  and  ventral    band.     (4)    The 


7*» 


Fig.  326. — Ancylostoma  duodenale :  diagrammatic  representation  of  excretory  system 
ex.p.,  excretory  pore;  e.c.g.,  excretory  cervical  gland  ;  Ex.  ves.,  excretory  vesicle  in  B.c.y 
bridge  cell,  which  is  connected  with  c.g.^  cervical  gland,  and  /./.,  lateral  lines;  ceph.g.^ 
cephalic  gland  ;  l.ex.c,  lateral  excretory  canal  passing  into  the  bridge  cell ;  /./.,  lateral  line 
containing  excretory  canal  and  cephalic  gland;  c.p.,  cervical  papilla;  w.,  nuclei  of  bridge 
cell.     (After  a  drawing  of  Looss.) 


ventral  band  is  well  developed  caudally,  forming  a  large  pad  dorsal 
to  the  cloaca,  "pulvillus  post-analis." 

The  bursal  rays  are  outgrowths  of  the  lateral  lines.  Beside  this 
^'  pulp  "  they  contain  a  nerve,  and  at  their  bases  complex  muscles. 

The  Bursa  is  closed  on  all  sides  with  a  short  median  (ventral)  lobe, 
which  may  be  tucked  inwards.  It  is  an  outgrowth  of  the  inner  layer 
of  the  skin  pushing  the  outer  layer  before  it,  so  that  it  consists  of  three 
layers,  not  four,  as  it  would  be  if  it  were  a  fold.  The  bursa  is  twice 
as  broad  as  long.  It  is  supported  by  a  variety  of  rays,  the  arrange- 
ment  of   which    is    best   followed   from   the  figure   (fig.   327).     The 


ANCYLOSTOMA   DUODENALE  449 

different  terminology  for  these  rays  as  used  by  various  authors 
should  be  noted :  Ventral  =  anterior  ;  externo-lateral  =:  antero- 
external ;  medio-lateral  +  postero-lateral  or  antero-median  +  postero- 
median =  median  (doubled);  externo  -  dorsal  =  postero  -  external ; 
dorsal  =  posterior.  All  the  rays  end  in  tactile  papillae,  seven,  on  each 
side  ;  the  postero-external  and  antero-external  on  the  outer  surface  of 
the  bursa,  the  five  others  on  the  inner  surface/  Of  the  six  terminal 
digitations  of  the  dorsal  ray,  only  the  external  two  contain  tactile 
papillae. 

In  the  male  there  are  prebursal  papillae  and  minute  caudal  papillae 
in  the  female. 

In  the  female  the  inner  layer  of  the  cuticle  projects  at  the  posterior 
end  as  a  sharp  spike,  20  /x  long,  which  may  sometimes  be  broken  off. 

Ovaries. — The  anterior  tube  runs  from  the  cephalic  to  the  posterior 
end  and  back  again.     The  posterior  tube  begins  anteriorly,  runs  to  the 


Ci/e 


Fig.  327. — Ancylostoina  duodenale :  bursa  enlarged.  Ca,  anterior  rpy 
cleft  ;  cle,  antero-external ;  els,  antero-median  ;  dp,  postero-median  ;  Cde, 
postero  external  ;  Cd,  posterior  bifurcated,  each  bifurcation  tridigitate. 
After  Railliet.) 

posterior  end  of  the  body,  and  then  back  to  the  cephalic  end,  form- 
ing a  vulval  loop  before  ending.  The  ovaries  on  the  whole  run  in 
oblique  coils.  The  uterus  is  the  thicker  portion  of  the  tube,  5  mm. 
long.  A  short  tube  connecting  the  ovary  and  uterus  is  the  oviduct. 
The  two  uteri  unite  to  form  a  single  duct,  the  vagina,  opening  i  mm. 
behind  the  middle  line.  The  portion  of  the  uterus  next  to  the  oviduct 
functions  as  a  seminal  receptacle,  whereas  the  part  next  the  vagina 
functions  as  an  ovejector. 

Testis. — The  blind  end  begins  a  little  behind  the  beginning  of 
the  cement  gland.  The  transverse  coils  occupy  the  middle  third  of 
the  body.  About  the  middle  of  the  body  it  passes  into  the  spindle- 
shaped  seminal  vesicle,  which,  with  the  spicular  canal  and  rectum, 
opens  into  the  cloaca.  An  anterior  longitudinal  coil  pushing  in 
between  the  cervical  glands  is  characteristic  of  Ancylostoma.  The 
cement  gland  surrounds  the  ejaculatory  duct  for  practically  its  whole 
course,  and  it  occupies  nearly  the  posterior  half  of   the  body  and 

1  This  also  occurs  in  other  Strongylida,  e.g.,  in  the  genus  Strongylus  (Syn.  :  Scleroatomum). 


450 


THE    ANIMAL   PARASITES   OF   MAN 


secretes   a    brown    or  black  cement.     The   spermatozoa   are    curved 
rods  about  2  fi  long. 

Spicules  are  2  mm.  long,  ending  in  a  fine  point.  They  are  moved 
by  exsertor  and  retractor  muscles.  At  first  they  lie  free  in  the  body 
cavity;  next  in  a  groove  in  the  dorsal  wall  of  the  cloaca;  then  in  an 
isolated  canal,  and  finally  in  two  canals.  Anteriorly  each  has  two 
longitudinal  crests  on  its  inner  surface.  These  meet  the  correspond- 
ing crests  of  the  other  spicule,  and  so  form  a  canal  along  which  the 
sperm  passes  into  the  female.  The  gubernaculum  is  a  thickening 
of  the  dorsal  wall  of  the  cloaca.  It  is  not  a  free  piece,  but  is  moved 
by  various  muscles. 


Fk;.  328. — Ancylostoma  duodeiiale  :  bursa  of  male.  The  rays 
from  left  to  right  are:  (i)  anterior  clef  t ;  (2)  antero-external ;  (3) 
and  (4)  median  doubled,  i.e.,  antero-median  and  postero-median  ; 
(5)  postero-external  arising  from  a  common  trunk  with  the  posterior. 
X  c.  120.     (After  Looss.) 

Genital  Cone  is  a  prominence  on  the  floor  of  the  bursa  on  the 
ventral  side  of  the  body,  on  which  the  genito-anal  orifice  opens. 
The  cone  is  only  slightly  marked  in  Ancylostoma  duodenale,  but  is 
much  more  prominent  in  Necator  aniericanus.  ^ 

Distribution. — Africa,  Egypt,  Europe,,  Japan,  China  (mainly),  but  in 
association  with  Necator  aniericanus  in  Southern  States  of  America, 
British  India,  Assam,  Burma,  Hongkong,  Liberia,  Jamaica,  Martinique, 
Costa  Rica,  Colombia,  Antigua,  Guadeloupe. 

Habitat. — The  worms  live  in  the  jejunum,  less  frequently  in  the 
duodenum,  of  man  only. 

Food. — The  worms  feed  on  the  mucous  membrane  of  the  gut," 
attaching  themselves  to  the  base  of  the  villi,  sucking  these  in;  and 
when  these  are  destroyed  they  attack  further  the  submucosa.  As 
a  rule  the  worms  have  no  blood  in  the  gut,  but  in  their  attack  on 
the  submucosa  a  blood-vessel  may  be  eroded,  and  so  the  gut  of  the 
worm  filled  with  blood. 


ANCYLOSTOMA    DUODENALE 


451 


Development. — The  eggs  are  oval  with  broadly  rounded  poles, 
56yLt  to  61  fx  by  34//,  to  38  yu-.  In  fresh  faeces  they  contain  four 
granular  nucleated  segmentation  masses  of  the  ovum  (fig.  329) 
separated    by  a  clear  space   from  the  shell. 

Egg  of  Ancylostonie  appears  to  have  a  single  contour.  Under  high 
powers  this  appears  double,  but  they  are  the  outer  and  inner  surface 
of  the  true  (chitinous)  egg-shell.  Internal  to  this  is  the  extremely 
delicate  yolk-envelope,  a  kind  of  skin  secreted  by  the  Qgg  cell  around 
itself  for  protection.  The  function  of  this  is  probably  to  absorb 
water  to  swell  and  burst  the  outer  chitinous  shell.  The  embryos 
when  hatched  are  termed  larvae. 

Embryos  which  are  ready  to  hatch  have  their  bodies  almost  free 
from  granules  ;  others,  though  apparently  mature,  that  have  granules 
will  not  hatch. ^ 

a  h  c  d 


Fig.  329. — Ancylostonia  auodenale :  eggs  in  different  stages  of  develop- 
ment, a  to  c,  in  fresh  faeces  ;  d,  containing  a  larva,  only  in  old  faeces. 
X  336.      (After  Looss.) 

Larva. — Stage  I :  Average  length,  25  mm.  Maximum  thickness 
in  oesophageal  region,  17  fi.  Head  end  fairly  blunt,  from  behind  the 
anus  (the  tail)  tapering  in  an  uniform  manner.  Buccal  cavity  is 
characteristic,  10  /jl  to  12  fi  by  1  fi  to  S  /jl,  longer  and  narrower  than  the 
corresponding  larvae  of  Strongyloides  stercoralis.  CEsophagus  "  rhab- 
ditic  "  in  character,  i.e.y  it  has  three  sections,  but  they  are  not  so  clearly 
marked  off  as  in  larvae  of  the  genus  Rhabditis.  The  posterior  bulb  has 
a  Y-shaped  valve,  the  function  of  which,  according  to  Looss,  is  to 
prevent  regurgitation  of  food.  The  granules  of  the  gut  serve  as  a 
reserve  of  food,  and  are  used  up  if  the  larvae  are  starved.  The  genital 
rudiment   consists   of   two   cells   half-way  between   the   end   of   the 

'  Table  of  Differences  between  Larv^  of  A.  duodenale  and  S.  stercoralis. 


(i)  Vestibulum  oris 

(2)  Genital  rudiment 

(3)  Thickness 

(4)  CEsophagus    ... 

(5)  Tail     

(6)  Motion 

(7)  Gut      


A.  duodenale 


i'8  fi  broad 

3  fi  to  5  /i  long 

Thicker 

One-fourth  body  length 

Pointed 

Less  active  than 

Soon  fills  with  dark  granules 


S.  stercoralis 


25  /*  to  33  /t 

Half  body  length 
Two  fine  points 


Rhabditiform. 


Filariform. 


29 


452 


THE   ANIMAL  PARASITES   OF  MAN 


oesophagus  and  the  anus  in  the  mid-ventral  Hne.  The  larva  lives 
on  f^cal  matter  and  grows  to  about  0-4  mm.,  then  moult^  I  takes 
place  in  two  days  or  more,  the  skin  being  ruptured  by  the  activity  of 

the  larva. 

Stage  II :   The  larva  is  now  in  this  stage,  which  does  not  differ 
much  from  the  previous  one.      It  grows  to  0-5  mm.     The  mouth 


Fig.  330. — Ancylostoma  duodenale  larva  on  fourth  day  of  culture  on  right 
Strongyloides  stercoralis  larva  on  left.     (After  Leichtenstern.) 


Opening  closes.  The  oesophagus  elongates,  becoming  cylindrical  or 
"  filariform "  ;  a  new  skin  is  formed  underneath  the  old  one,  and  in 
about  a  week  moult  II  takes  place. 

Stage  III  :  The  mature  larva  remains  enclosed  in  the  old  skin. 
Its  movements  are  now  much  more  active  and  of  a  boring  character. 
Length  is  now  o'6  mm.  This  mature  stage  has  been  erroneously 
called  the  encysted  larva,  because  there  is  no  cyst  secreted  from  its 

'  Moults  take  place  by  the  formation  of  a  new  skin  below  the  old  one,  the  two  being  in 
close  apposition  at  first. 


ANCYLOSTOMA   DUODENALE 


453 


surface  by  the  larva,  but  it  is  simply  the  old  skin,  which  is  not  cast  off, 
but  is  retained  for  purposes  of  protection,  as  the  larva  is  free  living, 
but  casts  it  as  soon  as  it  assumes  parasitic  life  again.  From  the  egg 
to  this  mature  stage  is  thus  six  to  ten  days. 


Fig.  331. — Ancylostoma  duodenale:  left,  four  days  after  transmission  into 
dog,  190/1  ;  in  the  centre,  at  the  commencement  of  the  second  stage  of  develop- 
ment (five  to  six  days),  105/ 1  ;  on  the  right,  fourteen  to  fifteen  days  after  trans- 
mission.    42/1.     (After  Looss.) 

Bionomics  of  Development. — Air  :  Eggs  can  develop  when  shut 
off  from  the  air  for  a  ''  comparatively  long  "  time. 

Temperature  :  Hatching  takes  from  eight  hours  upwards.  Eggs 
develop  best  at  25°  to  30°  C,  but  will  not  develop  below  8°  to  10°  C. 
The  larvse,  however,  will  stand  freezing. 


454  THE   ANIMAL   PARASITES   OF   MAN 

Moisture :  Eggs  and  larvae  do  not  live  long  under  water,  because 
they  suffocate  or  starve,  but  mature  larvae  will  live  for  months  (six  to 
twelve)  in  water ;  they  require  no  food — in  fact,  can  take  none  in — 
but  live  on  their  reserve  granules,  and  in  course  of  time  become  as 
clear  as  glass. 

Thiguiotropisni :  The  mature  larvae,  after  casting  their  skin,  will 
penetrate  pith,  climb  up  stems,  stalks,  etc.,  and  creep  into  any  pore. 

It  is  important  to  recognize  that  this  third  stage  of  the  mature 
larva  is  the  only  infective  one. 

Mode  of  Entry  into  the  Body. — Infection  is  effected  through  the 
mouth  (Leichtenstern  and  others),  and  also  through  the  skin,  as  was 
first  discovered  by  Looss  and  afterwards  confirmed  from  the  most 
diverse  quarters,  partly  in  the  case  of  Ancylostoma  duodenale, 
partly  in  that  of  A.  caninuni  in  dog,  man,  and  monkey.  The 
larvae  that  gain  access  to  the  intestine  partly  through  contaminated 
food,  or  through  unwashed  hands,  or  under  some  circumstances 
through  water,  first  throw  off  their  "  sheath  " — that  is,  they  complete 
moult  II.  Moult  III  takes  place  four  to  five  days  after  they  have 
reached  the  gut,  and  they  now  have  a  mouth  capsule  supplied  with 
four  small  teeth  arranged  crosswise,  enabling  them  to  fasten  on  to 
the  intestinal  epithelium,  upon  which  they  feed.  On  this  food  the 
worms  grow  in  four  to  six  days  to  3  to  5  mm.  in  length,  and  now 
moult  IV.  takes  place,  thus  attaining  their  definite  shape  and 
distinctive  character.  About  eight  days  later  the  sexual  organs 
commence  to  function  ;  at  this  time  the  first  copulation  should  be 
taking  place — it  will  later  be  frequently  repeated — and  a  few  days 
later  the  first  ova  are  laid,  first  in  less  and  later  in  larger  numbers,  so 
that  they  appear  in  the  faeces  about  four  to  five  weeks  after  the 
infection.^ 

Infection  by  the  Skin. — Mature  larvae,  which  are  placed  on  the 
skin  of  man  or  suitable  animals,  cast  their  ^'  sheath  "  and  bore  their 
way  through  delicate  fissures  either  horizontal  in  the  superficial 
scales  of  the  epidermis,  or  through  vertical  fissures  into  hair  follicles 
where  these  exist,  and  then  they  invade  the  cutis.  Now  according 
as  they  migrate  further  into  the  lymphatic  vessels  or  the  small  vesicles, 
the  final  path  to  the  gut  differs  to  some  extent.  The  blood  path 
leads  to  the  right  heart,  and  from  there  into  the  lungs  ;  here  the 
larvae  leave  the  blood  stream  and  enter  the  air  passages,  over  the 
mucosa  of  which  they  travel  further  headwards,  through  the  bronchi 
into  the  trachea  and  larynx,  and  from  hence  through  the  oesophagus 
to    the   stomach ;    in    some   cases   also   they   are    swallowed.      The 

'  From  the  number  of  eggs  present  in  a  given  quantity  of  faeces,  the  number  of  female 
Ancylostomes  present  in  the  gut  can  be  reckoned  by  a  formula  of  Leichtenstern's  (x  =  ^'L,  in 
which  a  signifies  the  number  of  eggs  counted  in  a  single  gramme  of  fseces). 


ANCYLOSTOMA   DUODEXALE  455 

lymphatic  path  leads  finally  also  into  the  blood  stream,  but  the 
lymphatic  glands  must  first  be  passed,  and  in  these  many  larvae  are 
retained  and  perish.  In  the  cutaneous  infection  seven  to  ten  weeks 
elapse  till  the  time  of  appearance  of  the  first  ova  in  the  faeces. 

The  penetration  of  the  skin  by  the  larvas  also  in  man  causes  reddening  and 
burning  at  the  affected  points,  and  this  is  followed  in  a  few  days  by  transitory  swell- 
ing in  the  subcutaneous  connective  tissue.  Skin  affections  can  also  be  set  up  by 
such  Ancylostoma  (and  Strongyloides)  larvae  as  do  not  gain  access  to  the  blood  or 
lymphatic  vessels  or  gut ;  such  larvae  apparently  wander  further  in  the  connective 
tissue,  and,  as  Looss  has  in  his  own  person  observed,  gain  access  to  the  cutis  at 
different  points,  thus  causing  progressive  swellings  (accompanied  by  intense  itching), 
which  cease  when  the  worm  again  penetrates  into  the  deep  tissues.  Skin  affections 
such  as  "  ground-itch  "  or  "  pani-ghao  "  occurring  in  the  tropics  and  only  attacking 
the  feet,  or  other  affections  (^.^.,  sump  bunches)  are  now  well  recognized  as  being 
due  to  the  invasion  of  Ancylostoma  larvae. 

Other  names  for  these  skin  affections  are  water-sore,  sore  feet  of  coolies,  maza- 
morra,  bunches,  botches,  quaddeln,  kratze,  ampoules,  gourmes,  taons,  pitirr. 
Whether  oral  or  dermal  infection  is  the  more  important  one  further  observation 
must  decide. 

The  duration  of  life  of  Ancylostotna  duodenale,  which  is  a  specific 
parasite  of  man  and  has  not  been  observed  in  other  mammals, 
amounts  to  about  five  years,  as  strayed  larvae  according  to  Looss 
wander  for  this  extent  of  time  in  the  body. 

Cultivation  of  Larvce. — (i)  Mix  the  faeces  (free  from  drugs  such  as 
salines  or  thymol)  with  animal  charcoal,  adding  water  if  necessary 
till  a  consistence  of  porridge  is  obtained.  If  the  stools  are  very  fluid, 
allow  to  sediment  first  and  pour  off  the  fluid.  The  best  charcoal  is 
that  made  from  bones,  and  should  not  have  an  acid  reaction.  Char- 
coal is  necessary  in  order  to  prevent  fermentation,  which  kills  the 
larvae.  Spread  in  layers  2  to  3  mm.  thick  in  Petri  dishes.  Incubate 
at  room  temperature.  To  extract  the  larvae  from  the  culture  allow 
the  surface  thoroughly  to  dry,  then  pour  on  water ;  the  larvae 
wander  out  and  are  poured  off  and  subsequently  further  purified  by 
sedimentation  or  filtering  through  blotting  paper,  the  larvae  passing 
through. 

(2)  A  funnel  is  plugged  with  cotton  wool,  then  filled  with  washed 
sand  to  within  a  centimetre  or  two  of  the  rim.  Stand  this  in  a  jar  of 
water  so  that  the  level  of  the  water  is  slightly  below  that  of  the  sand. 
On  the  surface  of  the  wet  sand  now  place  layers  of  blotting  paper, 
and  spread  the  faeces,  diluted  if  necessary,  on  this  in  layers  of  a  few 
millimetres  thick  {vide  p.  474). 

Detection  of  Eggs. — Vide  p.  473. 

Dermal  Infection  of  Dogs. — Infection  with  larvae  of  A.  caninum. 
In  two  hours  most  of  the  larvae  are  free  in  the  cutis  and  in  four  hours 
in  the  subcutaneous  tissue.  By  scraping  a  few  days  later  the  mucosa 
of  the  trachea  large  numbers  of  larvae  are  found  there. 


456 


THE   ANIMAL   PARASITES   OF   MAN 


Ancylostoma  ceylanicum,  Looss,  191 1. 
At  the  anterior  edge  of  mouth  capsule  one  large  tooth  ;   below  or 
behind  this  towards  the  middle  line  a  very  small  tooth,  the  tip  only 
of  which  is  seen.  Male  5  mm.  average.      Lobes  of  bursa  almost  as 

long  as  broad,  strongly  projecting 
towards  the  ventral  side.  Rays  short 
and  relatively  thick.     Female  7  mm. 

Habitat. — Intestine  civet  cat  {Viver- 
riciila  nmlacensis),  Ceylon,  and  man  in 
Bengal  according  to  Clayton-Lane. 

Other  species  are  :  A.  caninum 
(Ercolani),  in  cat  and  dog,  Europe 
and  Africa;  A.  inalayamun  (Ales- 
sandrini),  1905,  in  the  Malay  bear 
(Helaretos  lualayanus)  ;  A.  pluriden- 
tatmn    (Alessandrini),    1905,    in    Felis 


Fig.  332. — Ancylostof/ia  ceylanicum  : 
head  end,  two  teeth  on  each  side,  the 
inner  almost  concealed  by  the  outer. 
X  c.  200.  (After  Looss.) 


niitis,  Brazil. 


Ancylostoma  braziliense,  Gomez  de  Faria,  1910. 

In  cats  (and  dog),  Brazil.     Female  8*5  mm.,  male  7*5  mm.  long. 
Eggs  65  fi  by  32  /jL.  Leiper  considers  it  to  be  identical  with  A .  ceylanicum. 


Fig.  333.^ — Ancylostoma  braziliense:  bursa  of  male.     (After  Gomez  de  Faria.) 

Group.      Bunostomeae,  Railliet  and  Henry,  1909. 
Bursa  with  median  double,  postero-  and   postero-external  arising 
from  a   common   trunk,    posterior  bifurcated,    each    limb    bidigitate 
(fig-  336)-  Vulva  in  middle  of  body  or  a  little  in  front.    Uteri  divergent. 


NEGATOR 


457 


Contains  the  following  genera  :  (i)  Bunostomum  (=  Monodontus)  ; 
(2)  Necator ;   (3)  Bathmostomum  ;  (4)  Gaigeria. 

Genus.     Necator,  Stiles,  1903. 

Mouth  capsule  small,  narrowed  anteriorly  (ventrally)  by  chitinous 
plates,  as  in  Uncinaria.  On  each  side  of  the  base  of  the  dorsal  cone 
a  lateral  chitinous  plate  or  lancet  with  smooth  edge  (not  serrated), 
ventral  lancets  as  in  Ancylostoma.  No  ridges  on  outside  of  ventral 
wall.  Aperture  of  dorsal  oesophageal  gland  on  tip  of  a  cone  pro- 
jecting freely  into  the  buccal  capsule.  Bursa  closed.  Posterior  ray 
cleft  to  its  root. 

Necator  americanus,  Stiles,  1902. 
Syn.  :  N,  africanus^  Harrison,  1910. 

Male  8  mm.  long,  female  10  mm.  The  head  is  strongly  bent  dorsal- 
wards  so  that  almost  by  this  character  alone  it  can  be  distinguished 
horn  Ancylostoma  duodenale.     The  buccal  capsule  is  markedly  small — 


Fig.  334.  —Necator  americanus.  Showing  cutting  plates  and  the  projecting 
dorsal  ridge,  and  deep  in  the  cavity  the  edges  of  the  ventral  lancets,  x  c.  475. 
(After  Looss.) 

in  the  male,  0*093  by  0*084  mm.,  in  the  female  o'li  by  0*097  mm. 
There  are  no  teeth  anteriorly  on  the  ventral  side  of  the  capsule,  but 
instead  there  are  two  cutting  chitinous  plates,  the  anterior  portions 
of  which  are  prominent  and  angular,  and  meet  in  the  middle  line 
in  front.  Posteriorly  on  each  side  the  plate  projects  less,  while 
between  the  anterior  and  posterior  parts  there  is  a  deep  angle.  The 
inner  (posterior)  ventral  lancets  which  also  occur  in  A.  duodenale  are 
large,  and  project  far  into  the  lumen,  the  tips  of  these,  of  the  lateral 
lancets,  and  of  the  dorsal  cone  almost  meeting  in  the  centre  of  the 
lumen.  As  already  stated  in  the  definition  of  the  genus  Necator,  there 
are  also  lateral  lancets  which  start  from  the  base  of  the  dorsal  cone. 
This  dorsal  ridge,  or  rather  in  this  case  cone,  is  a  striking  object  in 


458 


THE   ANIMAL   PARASITES   OF  MAN 


the  mouth,  and  projects  right  out  into  the  cavity,  and  on  its  summit 
opens  the  dorsal  oesophageal  gland. 


Fig.  335. — Necator  americamis :  lateral  view,  showing  the  dorsal  ridge  per- 
forated by  the  duct  of  the  dorsal  oesophageal  gland,  the  lateral  lancet  and  ventral 
lancet  and  the  nerve  papillae,      x  c.  475.     (After  Looss.) 

The  bursa  is  about  as  long  as  broad,  but  has  the  lateral  lobes 
strikingly  lengthened,  giving  a  trilobed  appearance  (fig.  336),  but  as 

in  Ancylostoma  diiodenale  it 
is  closed  on  the  ventral  side. 
The  distribution  of  the  rays 
is  best  understood  from  the 
figure.  The  genital  aperture 
lies  on  a  marked  conical  pro- 
tuberance ;  the  cement  gland 
is  bilobed  in  transverse  section. 
In  the  female  the  opening  of 
the  vulva  is  in  front  of  the 
middle  line,  in  A.  duodenale  it 
is  behind. 

The  spicules,  0*92  mm.  long 
are  hooked  at  the  extremity. 

Eggs  more  pointed  at  the 
poles  than  those  of  A.  duoden- 
ale, 6^  Ijuio  ^2  fi  by  36  yLfc,  so  that 
it  may  not  be  possible  to  dis- 
tinguish single  eggs  owing  to 
individual  variations,  yet  on 
comparing  a  number  they  can 
be  distinguished. 


Fig.  336. — Necator  americanus  :  bursa  of  male. 
The  rays  from  right  (top)  to  left  are:  (i)  pos- 
terior, (2)  postero-external,  (3)  and  (4)  median 
doubled,  i.e.,  postero-median  and  antero-median, 
(5)  antero-external,  (6)  anterior  (cleft),  and  above 
it  on  left  a  pre-bursal  ray.  x  c.  120.  (After 
Looss. ) 


SYNGAME^  459 

Geographical  Distribution. — Brazil,  Porto  Rico,  Cuba,  Central 
Africa,  East  Africa,  Victoria  Nyanza,  Gold  Coast,  Uganda,  North- 
Western  Rhodesia,  Ceylon,  Mysore.  For  other  localities  where 
A.   duodenale  is  also  found  see    p.  450. 

Habitat. — In  small  intestine  of  man  and  gorilla  {Troglodytes gorilla). 

Necator  exilidens,  Cummins,  1912. 

Syn.  :  A^.  africanus,  Looss,   1911. 

Male  7  mm.,  female  9  mm.  long.  The  edges  of  the  cutting  plates 
are  rounded,  not  angular,  and  do  not  meet  in  the  middle  line.  Inner 
(posterior)  ventral  lancets  very  small.  Lateral  lobes  of  bursa  broader 
than  long.      Rays  thick  and  plump. 

Habitat. — In  the  chimpanzee  {Anthropopithcciis  troglodytes), 

Ancylostomiasis. 
Morbid  Anatomy.  —  Organs  pale  and  bloodless.  Abdominal 
organs  sodden,  and  there  is  fluid  in  the  serous  cavities.  Lungs: 
oedema.  Kidneys  :  fatty  changes,  especially  large  pale  kidney. 
Liver  and  heart  also  show  fatty  changes— there  is  much  haemo- 
siderin  in  the  liver  cells.  Blood  :  early  stages,  a  leucocytosis  20,000 
upwards,  and  eosinophilia  50  per  cent.  Later,  anaemia  (hydraemia). 
The  number  of  worms  found  varies  from  ten  to  1,000.  They  are  rare 
in  the  duodenum,  but  occur  as  far  as  6  ft.  from  the  pylorus. 

Group.     Syngameae,  Railliet  and  Henry,   1909. 

Bursa  with  anterior  and  median  ray  cleft ;  antero-external,  close 
to  median;  postero-external,  arising  separately  from  posterior;  pos- 
terior bifurcate  to  base,  each  branch  bifurcate  or  trifurcate.  Vulva  in 
the  anterior  fourth  of  body.     Uteri  divergent. 

Genus.      Syngamus,  von  Siebold,  1836. 

Head  thickened,  not  tapering  ;  broad  mouth  with  gaping  buccal 
capsule. 

Male  and  female  often  in  permanent  copula. 

Parasitic  in  respiratory  passages  of  birds  and  mammals. 

Habitat. — S.  trachealis  in  poultry  ;  S.  bronchialis  in  goose ;  S.  laryn- 
getis  in  cattle ;  S.  vasicola  in  goats,  etc. 

Syngamus  kingi,  Leiper,  1913. 
Buccal  capsules  of  male  and  female  on  same  level.  In  S.  trachealis 
and  S.  laryiigeiis,  that  of  male  in  front  of  that  of  female.  In  S.  dispar, 
that  of  male  behind  that  of  female.  QCsophagus  of  male  one-sixth, 
that  of  female  one-ninth  of  total  length.  Mouth  capsule  in  male 
and  female  terminal ;  it  is  dorsal  in  S.  trachealis  and  in  mammalian 


4t)0 


THE   ANIMAL   PARASITES   OF  MAN 


Species.  Tail  of  female  bluntly  pointed.  Ovary  reaches  to  anus. 
Excretory  pore  opposite  the  middle  of  the  bulb  of  oesophagus.  In 
S.  trachealis  it  is  opposite  the  oesophageal  valves. 

Habitat — Found   in   sputum    of   patient   by  King   in    St.    Lucia. 
Normal  host  probably  a  carnivore. 


Fig.   337. — Syngamus  kingi  :  anterior 
end  of  male.     (After  Leiper.) 


Fig.  338. — Syngamus  kingi  :  anterior 
end  of  female.    (After  Leiper.) 


Family.     Physalopteridae. 
Genus.     Physaloptera,  Rudolphi,  1819. 

Mouth  surrounded  by  two  large  lateral  lips  bounded  posteriorly 
by  a  cuticular  band  projecting  anteriorly,  forming  a  collar.  Each  lip 
bears  anteriorly  and  inwardly  a  cuticular  appendage,  the  external  tooth. 
Immediately  below  and  internal  to  the  external  teeth  the  internal  teeth, 
one  on  each  lip.  Each  lip  bears  two  large  submedian  papillae.  Tail 
of  male  with  four  pairs  of  pedunculated  papillae  in  a  row  on  each 
side  external  to  the  six  pairs  of  unpedunculated  papillae.  Spicules 
unequal.  Vulva  in  the  anterior  region  of  the  body.  Eggs  with  a 
characteristic  thick  smooth  shell. 

Parasitic  in  the  intestine,  more  especially  the  stomach,  of  mammals 
(twenty  species),  birds  (twelve  species),  reptiles  (fourteen  species). 


ASCARID^ 


461 


Physaloptera  caucasica,  v.  Linstow,  1902. 

The  male  measures  14*2  mm.  in  length  and  071  mm.  in  breadth  ; 

the  bursa  is  broad,  rounded  off  in  front  and  narrower  at  the  back ; 

the   right    spicule    measures    0*62    mm.    in    length,    the   left   spicule 

1*76  mm.  ;  there  are  two  papillae  in 
front  of  the  orifice  of  the  cloaca,  four 
behind  it  and  six  unpedunculated  on 
the  tail.  The  female  measures  27  mm. 
in  length,  1*14  mm.  in  breadth  ;  the 
caudal  extremity  is  rounded  off ;  the 
vulva  is  on  the  border  of  the  first 
and  second  sixth  of  the  length  of  the 
body;  the  eggs  have  thick  shells, 
and  measure  57  //,  by  39  //,.  It  has 
hitherto  only  been  observed  once, 
by  Menetries  in  the  intestine  of  man 
(Caucasus). 


Fig.  339. — Bursa  oi  Synga??ius  trachealis. 
a.,  anterior  ray  cleft  ;  a.e. ,  antero-external ; 
w.«.,antero-me(lian;  w./.,postero-median ; 
p.e.,  postero-external ;  /.,  one  branch  of 
posterior  (trifurcate).     (Stephens.) 


Physaloptera  mordens,  Leiper, 
1907. 

Large  worms  resembling  an  im- 
mature Ascaris  lumhricoides. 
The  inner  lancet-shaped  teeth  have  a  sharp  cutting  edge  towards 
the  lumen.     Below  each  is  a  cuticular  boss  projecting  into  the  mouth 

(fig-  340)- 

Male  30  to  50  mm.,  bursa  with  four  pairs  of  pedunculated  papillae, 
the  second  and  third  lying  external  to  the  first  and  fourth  on  each  side. 
Spicules  unequal,  one  slender  (4-6  mm.),  the  other  stouter  (6  mm.). 

Female  40  to  55  mm.  Tail  sharp.  Vulva  opens  between  the 
anterior  fourth  and  fifth  of  the  body.  Eggs  43*6//,  by  35-3  //,  with  a 
thick  smooth  shell. 

Habitat. — (Esophagus,  stomach,  small  intestine  of  man  (several 
cases).     Nyasaland  and  Portuguese  East  Africa. 


Family.    Ascaridae,  Cobbold,  1864. 

Sub-family.     Ascarinae. 

Without   oesophageal    or   intestinal    diverticula ;   spicules  without 
flanges. 

Genus.     Ascaris,   L.,  1758. 

Intermediate  lips  and  auricles  absent.    Lips  edged  with  fine  teeth.    Lips  triangular 
in  cross  section.     Not  grooved  on  internal  surface. 


46: 


THE    ANIMAL   PARASITES   OF   MAN 


.;  Slii.(». 


ves.Sim. 


Fig.  340. — Physaloptera  mordens,  Leiper,  1907.  (1)  adult  male  :  o.e.,  oesophagus;  ch.i.^ 
chyle  intestine;  t.c,  testicular  coils;  ves.  sem.,  vesicula  seminalis  ;  sp.i,  long  spicule;  sp.2, 
short  spicule;  B.,  bursa.  (2)  Mouth  parts:  r.,  cuticular  collar  embracing  the  two  lips 
posteriorly  ;  c.b.,  cuticular  bosses  guarding  the  mouth  laterally  ;  e.d.,  external  tooth ; 
i.d.,  internal  tooth  ;  stn.p.,  submedian  papillae.  (3)  egg  of  P.  caucasica.  (4)  egg  of 
P.  inordens.  (5)  bursa  enlarged  :  fed.p.,  pedunculated  papillae  ;  ses.p.,  sessile  papillae. 
(After  Leiper.) 


ASCARIS   LUMBKICOIDES 


Ascaris  lumbricoides,  L.,  1758. 


463 


The  colouring,  in  the  fresh  condition,  is  reddish-yellow  or  greyish- 
yellow  ;  the  body  is  of  an  elongated  spindle  shape.  The  oral  papillae 
are  finely  toothed.  The  dorsal  papilla  carries  two  sensory  papillae, 
the  two  ventral  papillae  each  one  sensory  papilla.  The  male  measures 
from  15  to  17  to  25  cm.  in  length,  and  about  3  mm.  in  diameter ; 
the  posterior  extremity  is  conical  and  bent  hook-like  ventrally ;  the 
spicules  measure  2  mm.  in  length,  are  curved,  and  somewhat  broadened 
at  their  free  end  ;  on  each  side  around  the  orifice  of  the  cloaca  there 
are  seventy  to  seventy-five  papillae,  of  which  seven  pairs  are  post-anal. 
The  testicular  tube  is  much  folded,  showing  through  the  body  integu- 
ment, and  is  about  eight  times  the  length  of  the  body.  The  female 
measures  20  to  25  to  40  cm.  in  length  and  about  5  mm.  in  diameteir ; 
the  posterior  extremity  is  conical  and  straight.  The  vulva  is  at  the 
junction  of  the  anterior  and  middle  thirds  of  the  body,  which,  at 


Fig.  341. — Ascaris  lumbricoides. 
extremity  of  the  male  with  the  spicules  pro- 
truding from  the  orifice  of  the  cloaca  {Sp.) ;  b, 
anterior  extremity  from  the  dorsal  surface,  the 
two  lobes  of  the  pulp  of  the  lip  separated  by  the 
"  saddle  "  ;  ^,  anterior  extremity  from  the  ventral 
surface;  /*.,  excretory  pore.  (From  Claus.) 


Fig.  342. — Ovum  of  Ascaris 
lumbricoides^  with  shell  and 
albuminous  envelope.     400/1. 


this  point,  has  a  slight  ring-like  constriction  ;  the  convoluted  ovaries 
measure  ten  times  the  length  of  the  body. 

The  ova  are  elliptical  with  a  thick  (4  />t)  transparent  shell  (fig.  342) 
and  an  external  albuminous  coating  which  forms  protuberances ;  the 
ova  measure  50  /i,  to  70  yu,  in  length,  40  ytt  to  50  //,  in  breadth ;  they  are 
deposited  before  segmentation  ;  the  albuminous  coating  is  stained 
yellow  by  the  colouring  matter  of  the  faeces,  but  is  sometimes  absent. 
The  egg  cell  is  unsegmented,  it  almost  completely  fills  the  shell,  and 
its  nucleus  is  concealed  by  the  large  amount  of  coarse  yolk  granules. 

Abnormal  or  unfertilized  eggs  also  occur  in  faeces.  They  are 
distinguished  by  their  elongated  form  (80  \ju  by  45  />t),  irregularly 
cylindrical,  its  contents  consisting  of  refractive  granules. 

Ascaris  lumhricoides  is  one  of  the  most  frequent  parasites  of  man ; 
it  is  distributed  all  over  the  inhabited  parts  of  the  world,  and  though 
it  is  particularly  frequent  in  the  warmer  regions,  yet  it  also  occurs 


464 


THE   ANIMAL   PARASITES   OF   MAN 


in  Finland,  Greenland,  etc.  In  temperate  climates  A.  lumhricoides 
occurs  most  frequently  in  young  children  ;  it  is,  moreover,  more 
common  amongst  country  dwellers  than  amongst  the  inhabitants  of 
towns,  but  is  not  lacking  in  infants,  adults  and  aged  persons.  As  a 
rule  only  a  few  specimens  are  present  in  the  intestine,  but  many  cases 
are  known  in  temperate  zones  in  which  several  hundreds  of  worms 
have  been  found  in  the  same  patient.  This  species  is  particularly 
numerous  in  the  negroes  of  Africa  and  America.  It  occurs  also  in 
the  monkey,  dog  and  pig  {1  A.  suilla). 

The  parasite  was  known  in  ancient  times  ;  the  Greeks  called 
it  eXjjLLv^  crrpoyyvXrj,  Plinius  termed  it  Tinea  rotunda,  ater  on 
it  was  named  Luntbricus  teres.  The  aaKapi^  of  the  Greeks  is  our 
Oxyuris. 

The  small  intestine  is  the  normal  habitat  of  Ascaris  lumhricoides ;  the  worms, 
however,  often  leave  this  part  of  the  intestine  and  wander  into  the  stomach,  whence 
they  are  frequently  evacuated  by  vomiting,  or  they  may  creep  through  the  oeso- 
phagus into  the  pharynx  and  crawl  out  through  the  nose  or  mouth  ;  very  rarely  they 
may  find  their  way  into  the  Eustachian  tube  or  into  the  naso-lachrymal  duct,  or  into 
the  excretory  ducts  of  the  liver  and  pancreas ;  exceptionally  they  may  gain  the 
trachea,  and  they  have  also  been  found  in  the  abdominal  cavity.  They  may  bore 
through  adhesions  between  the  intestinal  wall  and  the  omentum  (worm  abscess)  ; 
they  occasionally  penetrate  the  urinary  apparatus  and  are  passed  with  the  urine  ; 
in  febrile  diseases  ^. /«;«MV<9/^<?j-  usually  leaves  the  intestine  spontaneously.  It  is 
obvious  that  these  wanderings  may  be  accompanied  by  the  most  serious  symptoms, 
but  in  sensitive  persons  the  invasion  of  even  only  a  few  intestinal  Ascarides  gives 
rise  to  a  series  of  almost  inexplicable  symptoms .  (hysterical,  epileptiform  attacks, 
cerebral  congestion,  aphonia,  etc.),  which  cease_with  the  expulsion  of  the  worms,  so 
that  many  authors  are  driven  to  the  conclusion  _that  Jhese  Acarides  secrete  a  toxin. 
Fortunately,  the  presence  of  A.  lumhricoides  in  the  intestine  is  easily  demonstrated 
by  the  microscopical  examination  of  the  faeces. 

Development. — Several  authors  (Gros,  Schubart,  Richter,  Leuckart 
and  Davaine)  have  demonstrated  that  the  ova  of  Ascaris  develop 
in  water  or  moist  earth  after  a  long  period  of  incubation.  Freezing 
and  desiccation  (if  not  too  long)  do  not  injure  their  powers  of 
development ;  the  duration  of  the  development  depends  on  the 
degree  of  the  surrounding  temperature.  At  a  medium  temperature, 
after  a  varying  period  of  incubation,  it  takes  from  thirty  to  forty 
days  for  the  embryo  to  become  formed.  The  spirally  rolled  up 
embryo,  with  its  so-called  "  tooth,"  formed  by  three  papillae  close 
together,  never  leaves  the  egg-shell  in  the  open,  even  if  the  eggs  are 
kept  for  years  under  favourable  conditions.  Davaine  proved  that  the 
larvce  hatch  out  in  the  intestine  of  the  rat,  but  are  again  expelled  with 
the  faeces ;  he  therefore  concluded  that  the  hatching  likewise  takes 
place  within  the  intestine  of  man,  but  is  followed  by  the  invasion 
of  the  larvae.  In  the  meantime  Leuckart  had  sought  to  infect  himself 
by   swallowing   embryo-containing    eggs,    but   without    results  ;    he 


TOXASCARIS  465 

therefore  conjectured  that  there  must  be  an  intermediary  host,  and 
V.  Linstow  thought  he  had  found  it  in  myriapods  {JuUis  giittulatus). 
Subsequently,  Davaine's  opinion  proved  correct.  First  of  all  Grassi 
succeeded  in  infecting  himself  by  swallowing  100  embryo-containing 
eggs  of  Ascaris  Inmbricoides  ;  five  weeks  after  ingestion  the  worms  had 
attained  maturity  and  their  ova  appeared  in  the  faeces.  Calandruccio 
also  sought  to  infect  himself,  but  failed,  yet  he  succeeded  in  infecting 
a  little  boy  aged  7.  Lutz  also  reports  a  successful  experiment  which 
must  have  been  positive,  as  young  worms  5*5  to  18  mm.  long  were 
expelled.  Lutz  proved  that  the  eggs  lost  their  albuminous  sheJJ  by 
long  lying  in  water  and  then  died  when  introduced  into  the  stomach ; 
this  would  explain  the  failure  of  Leuckart's  experiment ;  in  moist  earth 
the  albuminous  shell  is  retained.  Finally,  Epstein  conducted  unim- 
peachable experiments  on  three  children  which  place  direct  infec- 
tion with  embryo-containing  eggs  beyond  doubt;  he,  moreover, 
proved  that  the  development  of  the  eggs  takes  place  more  rapidly  in 
the  faeces  when  there  is  free  admission  of  air,  sun,  and  a  sufficiency 
of  moisture. 

Accordingly,  infection  occurs  partly  through  water,  but  principally 
direct  from  the  soil. 

Ascaris,  sp. 

Wellmann  states  that  yet  another  species  of  Ascaris  in  man  occurs 
in  the  highlands  of  Angola  :  up  to  the  present  nothing  certain 
is  known  about  it  (Welland,  ^'Critical  Notes  on  Tropical  Diseases 
of  the  Angola  Highlands,"  New  York  Med.  Journ.  and  Philadelphia 
Med.  Journ.,  August  12  to  September  2,  1905.) 

Ascaris  texana,  Smith  et  Goeth,  1914. 
Female   alone   know^n  ;    58   to  60  mm.  and  upwards   in   length ; 
characterized  by  the  serration  of  the  anterior  border  of  the  lip  and  by 
the  appearance  of  interlabia.     Evacuated  by  a  white  settler  in  Texas. 
Position  of  this  worm  doubtful. 

Ascaris  maritima,   Leuckart,  1876. 

Only  one  immature  specimen,  a  female  (43  mm.  in  length  and 
I  mm.  in  breadth),  has  hitherto  been  described,  and  it  was  vomited  by 
a  child  in  North  Greenland  in  1865.  (R.  Leuckart,  '^  Die  menschlichen 
Parasiten,"  1876,  edition  2,  i,  p.  877.) 

Genus.     Toxascaris  (ro^ov,  an  arrow),  Leiper,  1907. 

Body  anteriorly  bent  dorsally,  cuticle  finely  striated.  CEsophagus 
without  a  distinct  bulb.  Tail  of  male  tapers  to  a  point.  Testis  in 
anterior  portion  of  posterior  half  of  body.  Vulva  about  middle  of 
body.     Eggs  oval  and  smooth. 


466  THE   ANIMAL   PARASITES   OF   MAN 

Toxascaris  limbata,  Railliet  and  Henry,  1911. 

Syn.  :  Lumbricus  cants,  Werner,  1782  ;  Ascarts  teres,  Goeze,  1782  ;  Ascaris  cati 
et  canicidce,  Schrank,  1788  ;  Ascaris  cams  et  felis,  Gmelin,  1789  ;  Ascaris  tricuspidata 
et  felis,  Bruguiere,  1791  ;  Ascaris  werneri,  Rud.,  1793;  Fusaria  mystax,  Zeder, 
1800  ;  Ascaris  7narginata  et  inystax,  Rud.,  1802  ;  Ascaris  alata,  Bellingham,  1839. 

Striations  6  //,  to  12  />«,  apart.  Cephalic  wings  long,  narrow,  semi- 
lanceolate.  Male,  4  to  6  cm.  Spicules,  1,002  y,  and  1,005  M- 
Female,  o'5  to  10  cm.  Eggs,  75  /x  to  85  /a,  shell  thick  and  smooth. 
Host :  dog,  occasionally  man. 


Fig.  343. — Ovum  Fig.  344. — Transverse  section  through  the  head 

of    Toxascaris     livi-  part  of  Belascaris  cati  from  the  cat,  with  the  lateral 

bata,  with  thin  albu-  wings.      In  addition,  one  n)ay  note  the  four  fields  of 

minous         envelope.  muscles,  the  longitudinal  lines  with  the  oesophagus 

Magnified.  in  the  centre.     Magnified.     (After  Leuckart.) 

Genus.     Belascaris  (/Se\o9,  an  arrow),  Leiper,   1907. 

Body  anteriorly  bent  ventrally,  cuticle  coarsely  striated. 
(Esophagus  with  a  distinct  bulb.  Tail  of  male  conical.  A  papillce- 
bearing  protuberance  behind  the  anus.  Testis  in  anterior  half  of  body. 
Vulva  in  anterior  part  of  body.     Eggs  corrugated. 

Belascaris  cati,  Schrank,  1788. 

Syn.  :  Belascaris  mystax,  Leiper,  1907  ;  Ascaris  mystax. 
Striations  12  /x  to  16  /i,  apart.     Cephalic  wings  lanceolate.     Male 
3  to  6  cm.    Spicules  17  to  1*9  mm.    Female  4  to  10  cm.      Eggs,  65  /t 
to  75  //<  in  diameter,  surface  finely  honeycombed.       Host  :  domestic 
cat,  and  man,  eight  or  nine  cases. 

Belascaris  marginata,  Rudolphi,  1802. 

Striations  16  /a  to  22  yu-  apart.  Cephalic  wings  long,  narrow,  semi- 
lanceolate.  Male,  5  to  10  cm.  Spicules,  750  /x  and  950  /x.  Female, 
9  to  18  cm.  Eggs,  75  /A  to  80  II.  Shell  finely  honeycombed.  Host  : 
dog. 

Genus.     Lagocheilascaris,  Leiper,  1909. 

Thick  lips  separated  by  a  furrow  from  the  body ;  between  the  lips 
small  intermediate  lips  without  "pulp."  The  cutting  angle  of  each 
lip  bifurcated.  Along  each  lateral  line  a  cuticular  wing  extending  the 
whole  length  of  the  body.     Eggs,  thick  shell  with  a  mosaic  pattern. 


OXYURID^  467 

Lagocheilascaris  minor,  Leiper,  1909. 

Male,  9  mm.,  tail  sharply  curved.  Spicules  colourless,  3*5  and 
4  mm.  long.  More  than  twenty-four  pairs  of  pre-anal  papillae,  at  least 
five  pairs  of  post-anal.  Female,  15  mm.  Straight  posteriorly.  Vulva 
6  mm.  from  head  with  two  lips.  Eggs,  65  />t  in  diameter.  Host  : 
man,  cutaneous  abscesses.     Trinidad. 

Family.     Oxyuridae. 

Genus.     Oxyuris,  Rudolphi,  1803. 

Mouth  unarmed.  The  three  labial  papillae  are  only  slightly  protuberant,  the 
oesophagus  is  long  and  presents  two  well-marked  bulbs.  The  vulva  is  in  the  anterior 
part  of  the  body. 

Oxyuris  vermicularis,   Linnaeus,  1767. 
Syn.  :  Ascaris  vermicularis^  L.  ;  Fusaria,  Zeder,  1803. 

Colour  white,  the  striated  cuticle  forms  projections  at  the  anterior 
end  w^hich  extend  some  distance  back  along  the  middle  of  the  ventral 
and  dorsal  surfaces  ;  the  longitudinal  lateral  flanges  of  the  skin  corre- 
sponding to  the  lateral  lines  are  well  seen  in  transverse  sections  ;  there 
are  three  small  retractile  labial  papillae  around  the  mouth.  The 
male  measures  3  to  5  mm.  in  length,  and  shortens  on  death;  the 
posterior  extremity  of  the  body  is  curved  ventrally  and  presents  six 
papillae.  Spicule  70/x,  long,  hook-like.  The  female  is  10 mm.  m  length 
and  0*6  mm.  in  diameter  ;  the  anus  is  about  2  mm.  in  front  of  the  tip  of 
the  tail ;  the  vulva  is  in  the  anterior  third  of  the  body ;  the  eggs  are 
oval,  asymmetrical,  with  double-contoured  shells,  and  measure  50  ^  to 
55//,  by  16  fjb  to  25//,;  they  are  deposited  with  clear,  non-granular 
tadpole-like  embryos  already  developed. 

Habitat. — Adults  in  large  intestine  of  man.  Young  forms  in  small 
intestine  and  often  in  the  appendix. 

The  worm  lives  in  the  lower  part  of  the  small  intestine,  caecum  and  vermiform 
appendix,  and  before  becoming  adult  undergoes  two  or  three  moults  (Heller). 
According  to  Wagener  the  worms  at  times  live  in  the  gut  wall,  giving  rise  to  calcareous 
nodules.  When  the  uterus  of  the  fertilized  females  begins  to  fill  with  eggs  they  leave 
the  caecum  and  travel  through  the  colon  to  the  rectum.  The  uterus  is  now  packed 
with  eggs  which  contain  a  tadpole-shaped  embryo.  Egg-laying  now  takes  place, 
partly  in  the  rectum,  partly  outside,  the  mode  of  exit  being  not  only  passive  through 
defsecation  but  also  an  active  one  on  the  part  of  the  worms  when  the  patient  is  in  bed. 
In  this  case  the  worms  crawl  out  of  the  anus,  producing  a  most  intolerable  itching  as 
they  scatter  their  eggs  between  the  nates  and  the  perinasum.  From  here  in  the  case 
of  girls  they  may  get  occasionally  into  the  vulva  and  vagina,  and  even  into  the 
oviducts  and  so  into  the  body  cavity.  The  worms  also  may  wander  through  the 
alimentary  canal  in  the  opposite  direction,  getting  out  occasionally  through  the 
mouth.  Recently  a  role  has  been  assigned  to  them,  as  to  other  gut  parasites,  in 
appendicitis  and  typhlitis. 

30 


468 


THE   ANIMAL   PARASITES   OF   MAN 


It  is  stated  that  the  males  die  after  fertilizing  the  females,  thus  explaining  why 
they  are  so  rarely  met  with  in  faeces  [but  it  is  probable  that  they  often  escape  notice 
from  their  small  size.— J.  W.  W.  S.]. 

Development. — The  eggs,  which  often  adhere  together,  contain  a 
tadpole-hke  embryo,  the  thin  tail  of  which  is  bent  upwards  ventrally ; 


Fig.  345--^, 
male,  and  B,  fe- 
male, of  Oxyuris 
vermicularis.  5/ 1 . 


Fig.  346. — On  the  left, 
female  ;  on  the  right,  male. 
A,  anus  ;  M,  mouth  ;  F, 
vulva.  Greatly  enlarged. 
(After  Glaus.) 


Fig.  347-—  Oxy- 
uris vermicularis  : 
egg  freshly  deposited, 
vk^ith  tadpole-like 
embryo,      x  640. 


Fig.  348. — Oxy- 
uris vermicularis  : 
egg  twelve  hours  after 
deposition,  with  ne- 
matode-like  embryo. 
X  640. 


the  embryo  in  a  short  time,  given  a  sufficiently  high  temperature,  passes 
into  a  second  folded  nematode-like  embryonal  stage,  lying  in  the  egg- 
shell, either  in  the  faeces,  with  which  also  numerous  females  pass  out, 
or  in  the  moisture  of  the  groove  between  the  buttocks,  and  they  there 
await  the  opportunity  of  being  reintroduced  into  man  per  os.     It  is 


MERMITHID^  469 

very  improbable  that  infection  takes  place  directly  in  the  large  intestine, 
as  is  occasionally  stated,  because  although  the  harbourers  of  Oxyuris 
are  frequently  liable  to  auto-infection,  this  takes  place  exclusively 
through  the  mouth,  and  is  conveyed  by  the  fingers,  on  which  the  ova 
of  Oxyuris,  and  occasionally  the  female  worms,  have  clung. 

The  opportunity  for  this  is  afforded  every  evening,  as  naturally  the  troublesome 
itching  caused  by  the  wandering  of  the  worms  is  met  by  scratching  and  rubbing  with 
the  fingers.  It  is  therefore  possible  that  the  eggs  may  even  thus  be  introduced  into 
the  nose,  where  the  young  Oxyuris  are  perhaps  hatched  out,  if  they  get  high  enough 
up  on  the  moist  pituitary  mucous  membrane.  As  a  matter  of  fact,  the  larvae  of 
Oxyuris  have  been  found  in  the  nose.  Moreover,  one  can  understand  that  the  eggs 
of  Oxyuris  are  transferred  from  person  to  person  by  the  hand,  directly  or  indirectly. 
This  again  explains  the  wholesale  infections  which  occur  in  collective  dwellings, 
after  a  person  harbouring  Oxyuris  has  been  admitted  into  boarding-houses,  etc. 
The  primary  infection  may  be  also  caused  in  other  ways — by  foods,  fruits,  vegetables 
and  other  articles  that  are  eaten  raw,  and  are  polluted  with  the  ova.  Perhaps  also 
flies  or  their  excrement  play  a  part  in  the  distribution  of  the  parasite,  similar  to  that 
demonstrated  by  Grassi  as  taking  place  in  the  spread  of  the  ova  of  Trichocephalus 
and  Taenia. 

The  assumption  of  a  direct  development  without  an  intermediary 
host  was  first  substantiated  by  Leuckart  by  experiments  on  himself 
and  three  of  his  students ;  about  fourteen  days  after  swallowing  the 
eggs  the  Oxyuris  has  attained  6  to  7  mm.  in  length  ;  Grassi,  and 
later  on  Calandruccio,  infected  themselves  by  swallowing  adult  female 
Oxyuris,  with  the  same  results.  Heller  found  worms  in  the  gut 
{appendix  vermiformis)  of  a  male  child  five  wrecks  old. 

Other  species  are  :  O.  co7npar  in  the  cat ;  O.  curvula  and  O.  mastigodes  in  horse, 
ass,  mule  ;  O.  ambigua  in  the  rabbit ;  O.  poculunt  in  the  horse  ;  O.  tenuicauda  in  the 
horse.  Many  species  occur  in  insects,  especially  in  Blattidce  and  Hydrophilidce 
(aquatic  beetles). 

Family.     Mermlthidae. 
Genus.     Mermis,  Dujardin,    1845. 
With  characters  of  the  family. 


Mermis  hominis  oris,  Leidy,    1850. 

Fourteen  centimetres  in  length,  o-i6  mm.  in  breadth;  mouth 
terminal ;  posterior  extremity  obtuse  and  provided  with  a  recurved 
hook  50  /^  long. 

The  parasite  was  "  obtained  from  the  mouth  of  a  child."  Stiles  considers  it  to 
be  probably  a  Mermis,  possibly  swallowed  in  an  apple. 


470  THE  ANIMAL   PARASITES   OP^   MAN 

Agamomermis,   Stiles,  1903. 
Group  name  for  immature  Mermithidce. 

Agamomermis  restiformis,  Leidy,  1880. 

This  worm  measures  65  cm.  in  length,  pointed  anteriorly,  the 
posterior  extremity  broadened  and  rounded  off  (1-5  mm.  in  breadth)  ; 
the  mouth  is  terminal,  without  lips.  Behind  the  mouth  six  papillae ; 
the  oesophagus  measures  i'i25  mm.  in  length  ;  the  intestine  appears 
to  terminate  blindly. 

This  parasite  was  obtained  in  West  Virginia  from  the  urethra  of  a  young  man, 
aged  20,  who  for  a  few  days  previous  to  expelling  the  worm  passed  turbid  and  bloody 
urine. 


TECHNIQUE  471 


TECHNIQUE. 

Preservation  and  Examination  of  Flukes. 

Fixation. — (Method  a.)  (i)  Place  the  flukes  in  a  test  tube  or  small  bottle 
a  quarter  full  of  normal  saHne.  Shake  the  contents  as  hard  as  possible  (the 
object  of  this  is  to  extend  the  flukes)  for  half  a  minute. 

(2)  Add  immediately  an  equal  bulk  of  saturated  aqueous  solution  of  corro- 
sive sublimate  and  shake  again  as  vigorously  as  possible  for  a  few  minutes. 

(3)  Transfer  when  convenient  to  70  per  cent,  alcohol.  (Before  staining 
and  mounting  remove  the  sublimate  with  tincture  of  iodine.) 

(Method  B.)  In  case  of  large  flukes,  e.g.^  Fasciola  hepatica,  Fasciolopsis 
buskij  compress  the  flukes  between  two  glass  slides  with  rubber  bands  or 
thread.    Fix  in  sublimate  or  in  absolute  alcohol,  or  in  10  per  cent,  formalin. 

(Method  c.)     Place  the  flukes  in  10  per  cent,  formalin  solution. 

Staining  is  successfully  effected  by  using  quite  dilute  solutions  of  carmine 
or  haematein  overnight.  This  is  far  preferable  to  using  strong  solutions, 
as  it  may  be  almost  impossible  to  remove  a  too  intense  stain.  Almost  any 
dilute  carmine  solution  suffices.  One  of  the  best  is  acetic-alum  carmine 
(boil  excess  of  carmine  in  a  saturated  aqueous  solution  of  potash-alum  for 
about  fifteen  minutes ;  add  glacial  acetic  acid  to  the  extent  of  10  per  cent. ; 
let  it  stand  for  a  week  ;  filter).  For  use,  dilute  about  thirty  times  with 
water.     Place  the  flukes  directly  in  the  stain.     Stain  overnight  or  longer. 

Differentiation, — In  order  to  get  the  sharpest  picture,  it  is  best  now  to 
differentiate  (but  this  may  often  be  omitted)  with  acid  alcohol  (70  per  cent, 
alcohol  100  parts,  HCl  5  drops).  Allow  to  act  from  one  to  twenty-four 
hours,  according  to  the  appearance  of  the  flukes.  Similarly,  in  staining  with 
hematoxylin  solution,  dilute  twenty  to  thirty  times  so  that  the  water  is  merely 
tinged  with  the  stain.  Differentiate  as  before.  After  staining,  dehydrate, 
clear,  and  mount  in  balsam  if  required. 

Clearing  and  Mounting. — (i)  Carbolic  acid  (carbolic  acid  94,  water  6)  is 
a  very  convenient  clearing  agent.  It  may  be  used  for  stained  or  unstained 
specimens.  It  will  clear  rapidly  without  previous  dehydration.  If  it  is 
required  to  mount  a  specimen  permanently,  transfer  from  carbolic  to 
alcohol,  then  cedar-wood  oil  (or  xylol,  etc.),  then  balsam. 

(2)  Creasnte. — Dehydrate  the  specimen,  stained  or  unstained,  transfer  to 
creasote.  If  it  is  desired  to  mount  permanently,  transfer  back  to  alcohol, 
then  cedar-wood  oil,  then  balsam'. 

(3)  Cedar-wood  Oil. — Preferable  to  xylol  or  oil  of  cloves.  Dehydrate 
the  specimen  in  alcohol.     To  mount  permanently,  transfer  to  balsam. 


472  THE   ANIMAL   PARASITES   OF   MAN 

(4)  Glycerine. — Vide  under  methods  of  preservation  of  ova;  to  mount 
permanently,  transfer  to  glycerine  jelly ;  subsequently  to  harden  the  jelly, 
expose  to  formalin  vapour. 

Of  these  media,  carbolic  acid  has  the  greatest  refractive  index  excepting 
that  of  balsam.  The  latter  may,  in  some  cases,  render  structures  too 
transparent,  and  it  may  be  advisable  to  use  only  glycerine  or  glycerine  jelly. 

Preservation  of  Ova  in  F^ces,  Urine,  Bile,  etc. 

Heat  some  70  per  cent,  alcohol  in  a  basin  to  about  60  to  70°  C.  (until 
bubbles  begin  to  appear).  Add  the  faeces,  etc.,  in  the  proportion  of  one 
part  to  about  nine  of  fixative  ;  keep  stirring.  Allow  the  sediment  to  settle. 
Transfer  to  a  bottle  with  some  fresh  70  per  cent,  alcohol. 

Transference  to  Glycerine. — Prepare  5  per  cent.,  10  per  cent.,  20  per  cent, 
solutions  of  glycerine  in  70  per  cent,  alcohol.  Pour  off  the  alcohol  in  the 
bottle  of  faeces,  etc.,  and  replace  by  5  per  cent,  glycerine  solution.  Allow 
to  stand  an  hour  or  so.  Then  in  the  same  way  replace  the  5  per  cent,  by 
a  10  per  cent,  glycerine,  and  finally  by  a  20  per  cent,  glycerine  solution. 
When  in  this  latter  expose  freely  to  the  air  (protecting  from  dust),  so 
as  to  allow  the  alcohol  and  water  to  evaporate.  Add  a  few  drops  of 
glycerine  from  time  to  time  till  eventually  the  ova  are  in  pure  glycerine. 
(In  a  very  moist  climate  it  may  be  necessary  to  use  lime  or  calcium 
chloride  to  dry  the  air.)  To  mount  permanently  transfer  some  of  the 
sediment  to  glycerine  jelly. 

Preservation  and  Examination  of  Cestodes. 

Fixation. — (i)  Saturated  aqueous  corrosive  sublimate. — Add  to  this  glacial 
acetic  acid  to  the  extent  of  i  per  cent.  (Note  this  fixative  will  dissolve 
the  "calcareous  corpuscles";  10  grammes  of  sublimate  to  160  c.c.  of 
water  will  give  a  saturated  solution.)  Warm  the  fixative  to  70°  to  80°  C. 
(Avoid  the  use  of  needles.)  Use  plenty  of  fixative.  Allow  to  act  for  a 
quarter  of  an  hour  or  so.  (a)  Transfer  to  70  per  cent,  alcohol.  (It  is 
advisable  to  remove  the  sublimate  by  the  use  of  Lugol's  solution,  or  a 
solution  containing  tincture  of  iodine,  adding  this  until  the  iodine  colour 
is  permanent.)     Or  (b)  transfer  for  preservation  to  10  per  cent,  formalin. 

Or  (2)  10  per  cent,  formalin. — In  order  to  prevent  contraction  it  is 
advisable  to  extend  the  tapeworm  and  keep  it  fixed  by  glass  plates,  or 
wind  the  worm  around  a  wide  glass  tube  or  bottle,  and  then  fix  it. 

Or  (3)  fix  in  hoi  alcohol. 

Staining. — As  under  flukes.  It  is  necessary  to  sacrifice  portions  of  the 
tapeworm  for  this  purpose,  cutting  out,  e.g.,  mature  segments,  so  as  to 
study  the  topography  of  the  genitaHa. 

Clearing. — As  under  flukes.  To  examine  the  hooks  satisfactorily  it  is 
best  to  cut  off"  the  head  with  a  sharp  knife  and  mount.  A  certain  amount 
of  pressure  is  then  advisable  in  order  to  view  the  hooks  completely  so  as 
to  measure  them. 


TECHNIQUE  473 


Preservation  of  Ova  in  F^ces,  etc. 
As  under  flukes. 

Preservation  and  Examination  of  Nematodes. 

Fixation. — (i)  Thoroughly  wash  the  worms  to  get  rid  of  mucus,  etc.,  by 
shaking  up  in  warm  saline  (or  water)  till  the  washings  are  clean.  Then 
transfer  to  70  per  cent,  alcohol  heated  to  about  70°  C.  It  is  absolutely 
necessary  to  use  hot  fixatives  in  order  to  extend  the  worms.  If  no  alcohol 
or  spirit  is  immediately  available,  drop  the  worms  into  hot  water,  or  saline, 
and  transfer  later  to  70  per  cent,  alcohol. 

(2)  Drop  into  hot  10  per  cent,  formalin. 

Cleaving. — (i)  Carbolic  acid,  vide  p.  471. 

(2)  Creasote,  vide  p.  471. 

(3)  Glycerine,  vide  p.  472. 

Staining. — In  case  of  quite  small  Nematodes,  e.g.,  Anguillulida,  carmine 
may  be  used,  but  as  a  rule  staining  is  not  advantageous. 

Rolling. — In  order  to  study  the  mouth  parts,  or  bursa,  etc.,  it  is  neces- 
sary to  place  the  worm  in  any  desired  position.  This  is  done  as  one  would 
roll  a  penholder  along  the  table  by  one's  finger  placed  on  top  of  it.  In  the 
case  of  a  worm,  one  edge  of  the  cover-glass  is  placed  over  the  worm,  the 
other  is  supported  by  a  strip  of  cardboard.  By  tapping  the  cover-glass 
the  worm  will  now  revolve  as  much  as  required  provided  it  is  round  and 
straight.  In  certain  cases  it  may  be  necessary  for  this  purpose  to  cut  off 
the  head  or  tail.     Roll  these  separately. 

When  a  suitable  position  is  got,  the  worm  may  be  fixed  in  this  position 
by  pressure  on  the  cover-glass,  so  as  slightly  to  flatten  it. 

Mounting  the  Head. — If  it  is  required  to  get  an  end  view,  it  is  necessary  to 
cut  off  the  head  transversely  as  near  the  end  as  possible,  and  then  mount. 

Detection  of  Eggs  (Bass  and  Hall). — Mix  the  faeces  thoroughly  with  ten 
times  the  volume  of  water.  Filter  through  gauze.  Centrifugalize  the 
filtrate.  Wash  the  sediment  and  centrifugalize.  Repeat  twice.  To  sedi- 
ment add  CaCl2  solution,  sp.  gr.  1250.  The  eggs  float  to  surface.  Pour 
off  surface  fluid.  Dilute  to  sp.  gr.  1050.  Centrifugalize.  Examine  sedi- 
ment, which  contains  practically  all  the  eggs  in  the  stool. 

Detection  of  Small  Nematodes. — Mix  the  faeces  thoroughly  with  water. 
Allow  to  settle  for  five  minutes.  Carefully  decant  off,  or  better,  syphon  off 
the  fluid.  Mix  the  sediment  again  with  water.  Allow  to  settle.  Remove 
the  fluid.  Repeat  several  times.  Examine  the  sediment  in  a  Petri  dish. 
As  the  fluid  is  poured  off,  the  worms  will  be  seen  collected  in  the  backwater. 
Remove  them  with  a  brush.     Fix  in  hot  70  per  cent,  alcohol. 


474  THE   ANIMAL   PARASITES   OF  MAN 

Cultivation  of  Larval   Forms    of   Ancylostoma  and  Strongyloides. 

A  modification  of  the  second  method  of  Looss  (p.  455)  is  that  of 
Fiilleborn.  A  glass  filter  funnel  is  lined  with  linen  or  with  cotton  wool 
dyed  black  with  iron-tannin.  On  this  is  placed  a  layer  of  sterile  sand,  and 
on  top  of  this  the  faeces.  The  whole  is  moistened.  The  larvae  hatch  out 
and  wander  through  the  meshes  of  the  wool,  appearing  on  the  edges  of  the 
same  as  white  threads  visible  to  the  naked  eye.  With  a  platinum  needle 
these  can  be  easily  removed.  The  glass  filter  can  be  placed  on  a  glass 
cylinder,  and  this  in  another  large  stoppered  cylinder  containing  caustic 
potash  solution  at  the  bottom,  so  that  any  larvae  escaping  from  the  funnel 
are  killed. 


ACANTHOCEPHALA  475 


D.  ACANTHOCEPHALA,  Rud. 

Gutless,  neraatode-like  worms  that  carry  at  their  anterior  end  a  retractile 
rostrum  beset  with  hooks.  In  their  adult  condition  they  only  live  in  vertebrate 
animals.     During  their  larval  stage  they  are  often  parasitic  in  invertebrate  animals. 

The  Acanthocephala  are  elongated  cylindrical  worms,  with  a  rounded  posterior 
end.  In  some  species  an  annulation  is  distinctly  recognizable  ;  they  are,  however, 
not  segmented.  The  size  varies  according  to  the  species,  between  about  $  to  lomm. 
and  40  to  50  cm. ;  in  general,  however,  there  is  a  preponderance  of  the  small  species. 
The  sexes  are  separate,  and  the  males  can  easily  be  distinguished  from  the  females 
without  examination  of  the  genitalia,  as  the  females  are  both  larger  and  thicker. 

The  body  wall  of  Echinorhynchus  is  limited  by  a  thin  cuticle,  which  is  attached 
inwardly  to  the  hypodermis.  In  only  exceptional  cases  a  syncytium  with  large 
nuclei,  even  in  the  adult  condition,  is  represented  by  the  hypodermis  ;  and  in  it  fibre 
systems,  the  elements  of  which  run  in  layers  in  various  directions,  appear,  and  it  is 
only  towards  the  interior  from  these  strata  of  fibres  that  the  nuclei  of  the  hypodermis 
are  found.  As  a  rule,  these  fibres,  at  all  events  the  radiary  fibres,  are  regarded  as 
muscles.  Hamann  desciibes  them  as  elastic  fibres,  which  lie  in  a  viscid  gelatinous 
connective  substance  (transformed  protoplasm  ?)  ;  a  lacune  system  filled  with  a 
granular  fluid,  the  central  part  of  which  are  two  longitudinal  lacunes  lying  at  the 
sides,  also  belongs  to  the  cutaneous  strata,  as  do  the  sc-called  lemnisci,  two  short, 
flat  organs  suspended  in  the  body  cavity,  and  the  pedicles  of  which  are  attached 
anteriorly  at  the  border  between  the  rostrum  and  body ;  their  structure  as  well  as 
their  origin  permit  them  to  be  traced  to  the  skin  (fig.  348A). 

Finally,  inwardly  below  the  skin  there  follows  a  layer  of  annular,  and  after  these 
a  layer  of  longitudinal  muscles,  the  structure  cells  of  which  remain  present  in  the 
residues,  carrying  nuclei.  The  motor  apparatus  of  the  rostrum,  the  sheath  of  the 
rostrum,  and  the  lemnisci  also  belong  to  the  muscular  system.  The  rostrum 
represents  a  finger-shaped  hollow  process  of  the  cutaneous  layer  ;  but,  according 
to  Hamann,  it  originates  from  the  entoderm  and  passes  through  the  skin  secondarily. 
It  is  covered  by  a  thin  cuticle,  and  as  a  rule  contains  a  large  number  of  regularly 
placed  chitinous  hooks  that  adjoin  a  granular  formation  tissue.  From  the  base  of 
the  rostrum  springs  a  tubular  hollow  muscle  extending  into  the  body  cavity ;  this 
is  the  RECEPTACULUM  PROBOSCIDIS,  from  the  base  of  which  again  bundles  of 
longitudinal  muscles  originate,  which  pass  along  its  axis  and  that  of  the  rostrum 
itself,  and  are  inserted  at  the  inner  surface  of  its  anterior  end  (RETRACTOR 
PROBOSCIDIS).  These  muscles  when  they  contract  invaginate  the  proboscis  and 
draw  it  into  the  receptaculum ;  when  reversed  they  act  again  as  PROTRUSOR 
PROBOSCIDIS.  The  whole  of  the  anterior  body,  however,  can  be  invaginaled,  and 
for  this  purpose  there  is  a  muscle  that  originates  from  the  body  wall  at  a  variable 
distance  back,  and  which  is  joined  to  the  receptaculum  (retractor  receptaculi)  ; 
there  is  also  a  bell-shaped  muscle  which  springs  from  the  body  wall  behind  the 
lemnisci  in  rings,  and  passes  forward  to  the  spot  of  attachment  of  the  lemnisci. 

The  nervous  system  consists  of  a  cluster  of  ganglia  situated  at  the  base  of  the 
rostrum,  from  which  three  nerves  pass  towards  the  front  and  two  towards  the  back. 
No  sensory  organs  are  known. 

The  excretory  organs,  according  to  Kaiser,  lie  at  the  upper  border  of  the  ductus 
ejaculatorius  in  the  male  and  at  the  so-called  bell  in  the  female.  Here  they 
30- 


476 


THE   ANIMAL   PARASITES   OF   MAN 


represent  the  long-known  villous  tufts,  placed  on  disc-like  cushions.  In  each  of  the 
cylindrical  villi — which  terminate  blindly  towards  the  body  cavity— there  lies  a 
cilium,  which  springs  from  the  membrane  lining  the  villus,  and  which  lies  in  a  space 

cavity  of  the  villus,  which  ultimately  proceeds  as  a 
little  canal.  There  are  three  canals  discharging  into 
the  uterus  that  serve  to  conduct  the  excretory  materials 
from  the  body  cavity  ;  special  glandular  cells  corre- 
sponding to  the  terminal  cells  of  the  Platyhelminlhs, 
at  the  commencement  of  the  system,  are  not  present 
in  the  Acanthocephala, 


\T. 


Fig.  348A.  —  The  male  of 
Echinorhynchus  augustatus. 
L.^  lemnisci ;  T.y  testicles;  A, 
prostatic  glands  ;  /*.?-.,  sheath  of 
proboscis,  with  ganglion  ;  R.r.^ 
retractor  of  sheath  of  proboscis. 
25/1. 


Sexual  Organs. 

{a)  Male  Organs. — The  greatest  part  of  the  male 
genital  apparatus  is  contained  in  a  muscular  sheath^— 
the  ligament — which  originates  at  the  posterior  end 
of  the  receptaculum  proboscidis,  passes  along  longi- 
tudinally through  the  body  cavity,  and  is  inserted  at 
the  posterior  end  of  the  worm.  The  two  oval  testicles 
usually  lie  one  behind  the  other  ;  their  vasa  efferentia 
unite  sooner  or  later  into  a  vas  deferens  which  passes 
backwards,  and  finally  terminates  in  the  penis  ;  the 
terminal  portion  of  the  conducting  apparatus  is  sur- 
rounded by  six  large  glandular  cells  (prostatic  glands) 


Fig.  348B. — ^Anterior  portion  of  the  female 
apparatus  of  Echinorhyvchus  acus.  On  the 
left  seen  from  behind,  on  the  right  seen  from 
the  front,  /^.inferior  orifice  of  the  bell;  B, 
bell;  IJg,  ligament;  M,  mouth  of  bell;  Ut, 
uterus.     Magnified.     (After  Wagener.) 


the  excretory  ducts  of  which  open  into  the  vas  deferens.  The  penis  itself  is  placed 
atthe  base  of  a  bell-shaped  invagination  of  the  posterior  end,  the  bursa,  which  is 
everted  during  copulation. 

{b)  Female  Organs. — There  are  only  two  ovaries  present  in  the  ligament  during 
the  larval  staged  During  the  course  of  growth  they  divide  into  accumulations  of 
cells  (placentula^,  loose  or  floating  ovaries),  which  finally  cause  the  ligament  to  burst 


ECHINORHYNCHUS   GIGAS  ^  477 

and  they  thus  attain  the  body  cavity.  Thence  a  peculiarly  constructed  apparatus 
finally  conveys  the  eggs  out.  This  apparatus  consists  of  the  uterine  bell  and  vagina, 
the  latter  discharging  at  the  posterior  extremity  of  the  body.  The  bell  is  a  muscular 
canal  provided  with  apertures  at  both  the  anterior  and  posterior  extremities.  Its 
interior  space  is  in  direct  communication  with  the  body  cavity,  and  the  anterior 
orifice  takes  up  all  materials  floating  in  the  cavity— egg-balls,  mature  and  immature 
eggs— and  pushes  these  further  backwards.  The  continuation  of  the  bell  lumen  is 
now  narrowed  by  a  number  of  large  cells  in  such  a  manner  that  only  bodies  of  a 
certain  form  can  pass  through  this  tract  and  attain  the  uteius;  everything  else  is 
conveyed  back  into  the  body  cavity  through  the  posterior  opening  of  the  bell. 

The  eggs  are  already  fertilized  in  the  body  cavity,  and  in  this  position  go  through 
their  development  to  the  formation  of  the  embryo.  Completely  developed  eggs  are 
surrounded  by  three  shells,  and  are  generally  fusiform.  The  eggs  agglomerate 
in  masses  in  the  uterus  until  they  are  finally  deposited  through  the  vagina  and  vulva. 
For  the  further  development,  the  transmission  of  the  eggs  into  an  intermediary  host — 
usually  a  crustacean  or  an  insect — is  necessary  ;  the  metamorphosis  is  very  compli- 
cated;  but  this  transmission  may  be  very  easily  effected  artificially  by  feeding  suitable 
crustaceans  {Asellus,  Gammarus,  etc.)  with  the  eggs  of  Aca7ithocephala  ;  this  being 
the  only  method  of  inducmg  the  larva  to  hatch  out  so  that  its  structure  may  be 
studied.  The  larva  appears  in  the  form  of  an  elongated,  somewhat  bent  body,  at  the 
stumpy  anterior  end  of  which  there  is  a  crown  of  hooks  or  spines,  whereas  the 
posterior  end  is  pointed.  Especial  retractors  draw  in  the  hook-beset  anterior  surface, 
and  an  elastic  cushion  beneath  them  jerks  them  forward  again  when  required.  In 
the  middle  of  the  body  a  roundish  heap  of  small  cells  is  seen,  from  which  the  entire 
body  of  the  Echinorhynchus  originates,  even  to  the  cutaneous  layer  ;  the  latter  is 
also  the  larval  skin  in  which  the  small  Echinorhynchus  gradually  grows.  The 
development  of  all  the  organs  takes  place  within  the  intermediary  host,  and  the 
parasite  only  needs  to  be  imported  into  the  terminal  host  to  attain  the  adult  stage  after 
a  certain  growth.     In  some  cases,  however,  a  second  intermediary  host  is  utilized. 

Species  of  Acanthocephala  only  occur  exceptionally  in  human  beings. 

Echinorhynchus  gigas,  Goeze,  1782. 
Syn.  :   Tcenia  hirudinacea^  Pallas,  1781. 

The  body  is  elongated,  gradually  decreasing  in  thickness  towards  the  back. 

The  rostrum  is  almost  spherical,  and  is  beset  with    five  or  six  rows  of  recurved 

hooks.      The  males  measure   10  to  15  cm.   in  length,  the  females  30  to  50  cm.; 

the  eggs  are  provided  with  three  shells,  of  which  the  middle  one  is  the  thickest. 

The  eggs  measure  o'o8  to  01    mm.    in  length.     The  giant 

Echinorhynchus  occurs  especially  in   the   intestinal  cana^ 

of  the  domestic  pig  ;  it   is  less  common  in  other  mammals. 

It  bores  deep  into  the  mucous  membrane  with    its  rostrum, 

and   causes   an   annular   proliferation    around   the   perforated 

spot ;  occasionally  also  it  causes  perforation  of  the  intestine. 

It  is  doubtful  whether  the  giant  Echinorhynchus   occurs 

in   man.     Leuckart  admitted  that   there  were  a  few  positive 

cases.      According    to    Lindemann,    Ech.   gigas    occurs    in 

human  beings  in  South  Russia,  and  its  presence  is  not  rare. 

This    statement,    however,    has    not     been    confirmed.      Its, 

^^^'   }^'^^'7~  ^^^      presence   in   man    is   by   no   means   impossible,  as   its  inter- 
01       Echinorhynchus      ^      ..  ,  ,  /  \     y,   c       1  njr  1  1     ^i    \ 

^igas.    300/1.    (After      mediary  host,  the   cankerworm,  or  cock-chater  {Meiolontha)^ 

Leuckart.)  is,  according  to  Schneider,  occasionally  eaten  raw  by  human 


47^  THE   ANIMAL   PARASITES   OF   MAN 

beings.  According  to  Kaiser,  the  golden  beetle  {Cetonia  aurata)  and,  according 
to  Stiles,  another  beetle  in  America  {Lachnosterna  arcuata)  are  also  intermediary 
hosts. 

Echinorhynchus  hominis,  Lambl,  1859. 

This  term  is  applied  to  an  Echinorhynchus  found  by  Lambl  in  the  intestine 
of  a  boy  who  had  died  of  leuccEmia;  the  worm  was  5*6  mm.  in  length,  and  the 
almost  spherical  head  was  beset  with  twelve  transverse  rows  of  hooks. 


Echinorhynchus  moniliformis,  Bremser,  1819. 


The  male  is  4  cm.  in  length,  the  female  8  cm.  long.  This  species  lives  in  the 
intestine  of  field-mice,  rats,  marmots  and  Myoxus  quercitius.  A  beetle  {Blaps 
mucronata)  is  the  intermediary  host. 

This   species   has   also   once   been    artifically  cultivated   in   man    (Grassi   and 


Calandruccio). 

a:f     i-^n  : 

:■  ":  -     e^f-.t.. 

.n\i    .  .:    -'«tiv. 

,^'^Aim':  '^  '..,-..,.: 

.J      .;..v.;    .r:.  *,.. 

!,,!.<..  •./    /■:•'.. '^  ! 

..^•..u  .-.. 

:  "  ■.-.■^4  .   \  :  .J, 

.  '          .:ru.i?i: 

on"i     ..              ?: 

c^'iU  L..-..   , 

•?^0!.-i.i,\'  / 

.U'-:.'[\.':i  'ej  ;■•  ^^;^ 

_.)■. -rtj^.v   y, 

' 

■  :-:;'nfjf'  C'l  ' 

h-.r; 


:!!j 


GORDIID^  479 


E.  GORDIIDAE. 

Very  long  thin  worms  similar  to  Filariae,  which,  in  their  adult  condition,  hve 
free  in  brooks,  pools  and  springs ;  the  mouth  and  the  commencement  of  the 
intestine  are  obhierated  ;  there  are  no  lateral  ridges,  and  the  muscular  system 
presents  a  structure  different  to  that  of  the  Nematoda .  The  posterior  end  of  the 
male  is  split,  and  spicules  are  lacking  ;  there  are  two  testicles.  In  both  sexes  the 
genitalia  discharge  through  the  terminal  gut. 

The  larvae,  which  carry  a  rostrum  beset  with  hooks,  force  themselves  into  the 
larvcC  of  water-hisects  :  more  rarely  they  invade  molluscs,  and  they  then  become 
encysted  within  the  body  of  the  host.  According  to  Villot,  at  least  a  part  of  them 
attain  the  intestine  of  fishes,  where  they  again  become  encysted,  and  after  a  period 
of  rest  they  travel  into  the  tissues  of  their  hosts,  and  finally  again  reach  the  exterior 
by  way  of  the  intestine,  where  they  then  become  adult.  In  most  cases,  however, 
the  gordius  larvas  are  taken  up  by  predacious  water  insects  ;  they  live  for  a  while 
in  the  body  cavity  of  these  insects,  undergo  a  metamorphosis,  and  finally  wander 
into  the  water. 

A  few  species  invade  man  accidentally  with  water,  in  which  case  they  are 
usually  vomited  up  : — 

Gordius  aquaticus^  Dujardin,  30  to  90  cm.  in  length  (Aldrovandi,  Degland, 
Siebold,  Patruban). 

Gordius  tolosanus^  Duj.,  11  to  13  cm.  in  length  (Fiori). 

Gordius  varius,  Leidy,  10  to  16  cm.,  female,  up  to  30  cm.  in  length  (Diesing).  . 

Gordius  chilensis^  Blanch.  (Guy).  Gordius  villoti^  Rosa  (Bercutti,  Camerano)  ; 
Gordius  tricuspidatus^  L.  Def.  (R.  Blanchard),  Gordius  violaceus^  Baird  (Topsent), 
and  Gordius  pustulosus^  Baird  (Parona).  j 

■J 

■     ^  ■  •  -i 


...  i    1  •     it.-.- 


4%> 


THE  ANIMAL  PARASITES  OF  MAN 


F.  HIRUDINEA   s.   DISCOPHORA     (Leech). 

The  Hirudinea^  which  have  been  appropriately  included  amongst  the  Annelida, 
differ  in  many  respects  from  the  typical  members  of  the  group;  their  body  is  long 
and  flat,  it  lacks  the  parapodia  that  are  characteristic  to  all  forms  of  Annelida  ;  but, 
on  the  other  hand,  possesses  a  terminal  posterior  sucker,  and  in  many  species  there 
is  also  an  anterior  sucker.  The  mouth  is  terminal  at  the  anterior  end,  the  anus  lies 
dorsally  above  the  posterior  sucker  (fig.  3480).  The  body  is 
segmented,  but  this  is  less  manifest  in  the  body  covering  than 
it  is  in  the  arrangement  of  the  internal  organs  ;  the  segmenta- 
tion, nevertheless,  is  also  indicated  exteriorly  by  the  appear- 
ance of  the  cutaneous  sensory  organs  which  correspond  to  the 
segments.  This  shows  what  the  condition  of  the  ganglia  in  the 
abdominal  ganglion  chain  has  taught  us,  that  the  anterior  and 
the  most  posterior  segments  are  considerably  abbreviated— a 
part  of  the  latter  taking  part  in  the  formation  of  the  suctorial 
organs.  In  a  great  many  species  the  skin  is  distinctly  annu- 
lated,  four  or  five  of  such  rings,  at  least  in  the  central  region 
of  the  body,  appearing  on  one  segment  of  the  body.  The 
condition  of  their  body  cavity  is  another  peculiarity  of  the 
Hirudineaj  it  is  narrowed  by  the  powerful  development  of  the 
connective  tissue  and  the  muscular  system  into  four  tubular 
sinuses,  which  have  the  appearance  of  blood-vessels.  There 
are  usually  one  dorsal  and  one  ventral  median  trunks,  as  well 
as  two  lateral  trunks  ;  in  addition,  a  particular  blood-vessel 
system  exists.  » 

The  skin  consists  of  a  very  thin  cuticle  that  is  cast  off  from 
time  to  time  ;  it  is  secreted  by  the  underlying  cylindrical  epi- 
thelium, which  contains  numerous  goblet  cells.  The  muscular 
system  is  strongly  developed  ;  it  consists  of  long  tubular  fibres, 
which  run  circularly,  longitudinally  and  in  the  dorso-ventral 
direction ;  the  muscular  system ,  is  subject  to  a  particular 
expansion  in  the  clinging  organs  and  at  the  commencement 
of  the  intestine.  On  the  whole,  the  alimentary  canal  represents 
a  tube  running  straight  from  the  mouth  to  the  anus,  which 
possesses  a  number  of  blind  sac-like  protuberances  at  the 
sides  varying  according  to  the  species.  The  most  anterior 
section,  the  pharynx,  in  the  leeches  with  maxillae  carries  three 
chitinous,  semicircular  plates  furnished  with  teeth  —the  jaws — 
which  serve  to  tear  up  the  epidermis  in  order  to  open  the 
blood-vessels  ;  in  the  leeches  with  rostra  a  long  protractile 
proboscis  rises  from  the  base  of  the  elongated  pharynx. 
Numerous  salivary  glands,  the  secretion  from  which  possesses 
toxic  properties,  discharge  into  the  pharynx.  The  oesophagus, 
which  follows  the  pharynx,  and  to  the  exterior  of  which 
numerous  radiary  muscles  are  fixed,  is  a  suctorial  organ  in 
its  entire  structure.  The  nutriment  in  the  larger  species  con- 
sists of  the  blood  of  vertebrate  animals,  in  smaller  species  and 
in  the  young  stages  the  food  consists  of  small  invertebrate 
animals. 


Fig.  348D.— The 
internal  organs  of 
the  leech.  The 
creature  has  been 
opened  from  the 
dorsal  surface,  and 
part  of  the  intestine 
has  been  removed. 
The  testicles,  with 
vas  deferens,  may  be 
seen  between  the 
blind  ducts  of  the 
intestine  ;  beyond 
these  on  either  side 
the  segmental  organs. 
The  female  genital 
organs  are  in  front 
of  the  most  anterior 
pair  of  testicles. 
(After  Kennel.) 


GNATHOBDELLID^  481 

The  NERVOUS  SYSTEM  exhibits  the  typical  structure  of  other  segmented  worms  ; 
the  sensory  organs  consist  of  the  previously  mentioned  goblet-shaped  cutaneous 
sensory  organs,  of  the  organs  of  taste,  and  of  eyes,  the  latter  frequently  being 
present  in  large  numbers. 

The  EXCRETORY  or  segmental  organs  exhibit  many  peculiarities,  which  cannot^ 
however,  be  detailed  here.  They  commence  with  funnels  in  the  lacunes  of  the  body 
cavity,  and  usually  discharge  on  the  ventral  surface.  ' 

Almost  all  the  Hirudmea  are  hermaphrodite  and  copulate  reciprocally.  The 
two  ovaries  are  very  small,  and  the  oviducts  that  proceed  from  them  soon  unite  into 
a  common  duct,  which  then  passes  into  the  uterus  and  discharges  through  the 
short  vagina  in  the  median  line  of  the  ventral  surface  behind  the  male, organs  into 
the  so-called  clitellar  region.  The  male  sexual  apparatus  consists  of  symmetrically 
arranged  testicles,  varying  in  number  according  to  the  species,  the  short  vasa 
efferentia  of  which,  one  by  one,  run  into  the  vas  deferens,  passing  towards  the  front 
on  each  side.  In  frqnt,  at  about  the  level,  or  a  little  in  front,  of  the  female. gejiitalia, 
the  two  vessels  pass  into  a  convoluted  mass  of  tubes  to  the  so-called  epididymis,  and 
then  discharge  into  the  single  protractile  penis  (fig.  348D). 

All  leeches  deposit  so-called  COCOONS.  These  are  small  barrel-shaped  or  pouch- 
like bodies,  which  are  surrounded  by  a  thicker  shell  and  contain  a  number  of  eggs 
in  a  large  mass  of  albumen  ;  the  albumen  originates  from  glands  of  the  generative 
organs,  the  shell  substance  from  cutaneous  glands  of  the  clitellar  region. 

Family.    Gnathobdellidae  (Leeches  with  Jaws).  ^  ,.; 

These  are  distinguished  by  the  possession  of  usually  three  jaws  in  the  pharynx"; 
the  body  consists  of  twenty-six  segments.  The  posterior  sucker  is  large  and  flat; 
the  anterior  sucker  is  smaller.  The  Hirudinea  have  five  pairs  of  eyes,  the  NephelititE 
have  four  pairs. 

Genus.     Hirudo,  L.,  1758. 

The  entire  body  consists  of  102  annulations,  five  appearing  on  one  segment  in 
the  central  region  of  the  body.  The  pharynx  has  three  semicircular  jaws,  the  arched 
border  of  which  is  beset  with  numerous  teeth  (50  to  100).  The  male  sexual  orifice 
lies  between  the  thirtieth  and  thirty-first  rings,  the  female  orifice  between  the  thirty - 
fifth  and  thirty-sixth.  There  are  numerous  species,  some  of  which  are  utilized  for 
medicinal  purposes.  ^       •  ^     ' 

J  ^  Hirudo  medicinalis,  L.,  1758. 

It  occurs   in   numerous   colour  varieties,  one  of  which   has   been    designated 

Hirudo  officinalis,    Moq.-Tandon.      Usually   the    dorsal    surface   is   greyish-green 

and    is    marked    with     six     rusty-red     longitudinal 

stripes.      The   ventral   surface   is   olive-green,   more 

or    less    spotted    with    black,    and    marked    at  the 

sides   with   a   black   longitudinal    line.     The    length 

averages    8    to    12   to    20   cm.      This   leech    lives    in 

swamps,  ponds  and  brooks,  overgrown  with  plants 

and    having     a     muddy    bed.      The     cocoons     are 

deposited    in    the    soil    at    the    sides.      Europe,   as 

well  as  North  Africa,  is  its  home.      At    the  present 

day  it   has   been   exterminated   from   most   parts  of 

Central    Europe,   but    it    is    still    very   common    in  a  1) 

Hungary.     Its    use    for    medicinal    purposes    is    well      ¥iG,   24^E.—JItni(fo  medicin- 

known.     A    large    leech    can    suck   about    15  grs.    of  ''''^-  ,  '^'     anterior     end       with 
:,,«>,,,  •     ,  f  ■  open  buccal  cauity,  with  the  jaws, 

blood,   and  about  the  same  amount   is  lost  through   j    ^^    ^j^g.   ^    ^^g  j^^  isolated. 

secondary  haeriiorrhage.  (After  Claus  ) 


482  THK    ANIMAL    PARASITES    OP^    MAN 

Hirudo  troctlna,  Johnston,  1816. 

Syn.  :  Hirudo  ittterrupta^  Moq.-Tandon,  1826. 

This  species  measures  8  to  10  cm.  in  length.  The  back  is  greenish,  with  six 
rows  of  black  spots  surrounded  by  red ;  the  lateral  borders  are  orange-coloured ; 
the  abdomen  spotted  or  unspotted.  Its  habitat  is  in  North  Africa  and  Sardinia.  It 
is  applied  medicinally  in  England,  Spain,  France,  Algeria,  etc. 

Genus.     Limnatis,  Moq.-Tandon,  1826. 

Nearly  related  to  Hirudo,  but  is  differentiated  by  a  longitudinal  groove  on  the 
inner  surface  of  the  upper  lip  of  the  anterior  sucker.  The  jaws  are  furnished  with 
over  100  very  sharp  toothlets. 

Limnatis  nilotica,  Savigny,  1820. 

Syn.:  Bdella  nilotica,  Sav. ;  L.  nilotica,  Moq.-Tandon;  Hcemopis  {vorax), 
Moq.-Tandon,  1826,/.  p.\  Hcefnopis  sanguisuga^  Moq.-Tandon,  1846  {nee  Hir. 
sanguis^  Bergm.,  1757). 

This  species  measures  8  to  10  cm.  in  length,  and  becomes  gradually  more 
pointed  towards  the  front ;  the  body  is  always  soft.  The  back  is  brown  or  greenish, 
and  has  usually  six  longitudinal  rows  (rarely  only  two  or  four)  of  black  dots.  The 
abdomen  is  dark  ;  but  numerous  colour  variations  occur. 

The  native  place  is  North  Africa,  especially  the  coastal  regions ;  it  is  also  found 
in  the  Canaries,  the  Azores,  Syria,  Armenia,  Turkestan,  perhaps  also  Southern 
Europe.  It  is  taken  into  the  mouth  with  drinking  water,  and  may  settle  in  the 
pharynx,  larynx,  oesophagus,  and  nasal  cavities  of  human  beings.  This  species  has 
also  been  observed  in  the  vagina  and  on  the  conjunctiva.  It  is  equally  fond  of 
attacking  domestic  animals. 

Hirudo  mysomelas  (Senegambia)  and  Hirudo  granulosa  (India)  are  placed  with 
this  genus,  and,  like  our  leech,  are  also  used  for  medicinal  purposes. 

J  Genus.     Haennadipsa,  Tennent,  1861. 

These  leeches  live  on  land,  and  measure  2  to  3  cm.  in  length.  About  a  dozen 
species  are  known.  They  are  a  veritable  scourge  to  persons  in  the  tropics  (Asia, 
South  America),  as  they  attack  them  to  suck  their  blood.  They  are  able  to  force 
their  way  even  under  close-fitting  garments,  so  that  it  is  difficult  to  protect  oneself 
from  their  assaults  {Hcemadipsa  ceylonica,  Bl.,  and  other  species). 

Family.     Rhynchobdeliidae  (Leeches  with  Rostrum). 

These  are  furnished  with  a  proboscis  in  lieu  of  the  jaws  ;  the  segment  consists 
of  three  annulations. 

Genus.     Haementaria,  de  FiHppi,  1849. 

Haennentaria  officinalis,  de  Fil. 
Inhabit  Mexico,  where  they  are  used  for  medicinal  purposes. 

Genus.     Placobdella,  R.  Blanch. 

Placobdella  catenigera,  Moq.-Tandon. 

Indigenous  to  South  Russia,  Hungary,  Italy  and  South  France.  It  is  a 
parasite  of  the  swamp  turtle,  but  frequently  attacks  human  beings. 


ARTHROPODA  483 


G.     ARTHROPODA     (Jointed-llmbed   Animals). 

BY 

FRED.  V.  THEOBALD,  M.A. 

Bilaterally  symmetrical  segmented  animals  which  are  covered  with  a  thick 
•cuticle  that  is  frequently  calcareous  {Crustacea)^  but  always  thinner  between  the 
segments  ;  they  carry  (primitively)  a  pair  of  jointed  appendages  on  every  segment.^ 
The  segments  of  the  body  are  uniform  in  certain  regions,  but  differ  from  those  of 
contiguous  regions,  so  that  it  is  easy  to  distinguish  three  parts  (head,  thorax  and 
abdomen),  each  composed  of  segments.  The  cephalic  segments  are  always  formed 
into  a  uniform  head,  the  segmentation  being  scarcely  recognizable  at  either  end  ; 
the  thoracic  segments  may  also  fuse,  or  part  or  all  of  them  may  coalesce  with  the 
head  ;  the  abdomen,  as  a  rule,  retains  its  segmentation,  but  this  may  possibly  also 
be  lost,  in  which  case  it  is  [sometimes]  united  to  the  cephalothorax.  The  structure 
of  the  three  regions  depends  mostly  on  the  varying  form  and  function  of  the 
appendages  :  those  on  the  head  are  primitively  locomotive  organs  (and  frequently 
are  still  so  in  the  early  stages),  but  they  become  transformed  into  feelers  and  mouth- 
parts  (mandibles,  maxillae)  ;  the  limbs  of  the  thorax,  however,  usually  retain  their 
ambulatory  functions,  as  frequently  do  those  of  the  abdomen  ;  sometimes,  however, 
the  abdominal  limbs  disappear,  entirely  or  partly;  in  the  latter  case  they  are  then 
utilized  for  other  purposes. 

In  their  organization  the  Arthropoda  approach  the  segmented  worms. 

The  Arthropoda  are  generally  divided  into  five  groups  {Crustacea^  Protracheaia^ 
Arachnoidea,  Myriapoda^  and  Insecta  or  Hexapoda\  of  which  only  the  Arachnoidea 
and  the  Hexapoda  interest  us  here. 

A.     ARACHNOIDEA  (Spiders,  Mites,  etc.). 

The  head  and  thorax  are  always  united  together ;  the  abdomen  is 
either  segmented  or  without  exterior  segmentation,  in  which  case  it  is 
united  with  the  cephalothorax.^     The  number  of  pairs  of  appendages 

'  [In  most  Arthropoda  the  skin  is  hardened  by  a  deposit  of  chitin  {Hexapoda,  etc). — 
F.  V.  T.] 

2  Parasitic  or  free-living  Crustaceans  may  now  and  then  invade  man  abnormally.  Thus, 
according  to  Betten,  C aligns  curtus  invade  the  cornea  (Betten,  R.  A.,  **  Par.  Crust,  as  a 
Foreign  Body  on  the  Cornea,"  Lancet,  1900,  i,  p.  1002;  and  Centralbl.  f.  Bakt.  u.  Far., 
xxix,  p.  506).  According  to  Laboulbene,  also  Gammanis  pulex  (Laboulbene,  A.,  "Obs. 
d'accid,  caus.  par  le  G.  ful.  apport.  avec  I'eau  de  boison  dans  I'estomac  d'un  homme,"  Bull. 
Acad,  med.,  1898,  p.  21). 

^  R.  Blanchard  has  compiled  thirty-five  cases  in  which  Myriapoda  have  been  observed  in 
the  intestine  as  well  as  in  the  nose  of  human  beings  ("Sur  le  pseudopar.  d.  myriap.  chez 
I'homme,"  Arch,  de  Par.,  1898,  i,  p.  452).  E.  Munoz  Ramos  reports  an  additional  case 
(ibid.,  p.  491).  A  few  years  ago  a  doctor  in  East  Prussia  sent  me  a  rain  worm  out  of  a  lady's 
nose  (of.  Hanan,  A.,  "  Wahrsch.  Pseudoparas.  v.  Schweiss  fliegenlarv.  u.  angebl.  Para?. 
V.  Regenwurmern  b.  ein  Hysterisch,"  Arch,  de  Par.,  1899,  ii,  p.  23). 

*  [This  is  only  so  in  the  Acarina  or  mites,  not  in  the  Aratieida  or  spiders. — F.  V.  T.] 


484  THE   ANIMAL   PARASITES   OF  MAN 

amount  to  six,  of  which  the  two  front  pairs,  the  chelicerae  and  the 
pedipalpi,  are  attached  to  the  head  region  and  the  four  remaining 
pairs  to  the  thoracic  region.^  The  abdomen  in  the  adult  condition 
has  no  appendages.  The  Arachnoids  are  air-breathers,  and  for  this 
purpose  are  either  provided  with  tracheae  or  with  so-called  lung-sacs, 
or  they  breathe  through  the  surface  of  the  body.  Some  aquatic  forms 
breathe  by  gills. 

There  are  eight  or  ten  orders  of  Arachnoids,^  of  which,  however, 
only  two,  the  Acarina  and  the  Linguatiilida,  have  to  be  considered 
here.^ 

Order.    Acarina  (Mites). 

Small  Arachnoids,  the  three  parts  of  the  body  of  which  are,  as  a  rule,  coalesced  ; 
it  is  only  rarely  that  a  faint  line  indicates  the  division  between  a  cephalothorax  and 
abdomen.  The  two  appendages  on  the  head  are  designed  for  biting  or  puncturing 
and  sucking,  and  vary  according  to  their  use.  The  chelicerae  *  are  fang-like  jaws 
or  puncturing  bristles  forming  a  kind  of  rostrum,  the  pedipalpi  are  claw-like  or  shear- 
shaped,  or  form  a  suctorial  proboscis.^  The  four  pairs  of  legs  are  usually  well 
developed,  more  rarely  they  are  rudimentary  or  have  partly  vanished  ;  many 
parasitic  forms  are  provided  with  pedunculated  suckers  [ambulacra — F.  V.  T.]. 
Respiratory  organs  (tracheal  tufts)  may  be  present  or  absent.  The  nervous  system 
is  reduced  to  a  minimum,  eyes  are  usually  lacking.  The  intestine,  situated  in  the 
central  part,  generally  has  three  blind  appendages  ;  the  anus  is  situated  on  the  venter 
above  the  posterior  end.  Sexes  separated  ;  nearly  all  the  species  deposit  eggs,  from 
which  six-legged  larvae  hatch.  The  Acarina  live  either  free  in  the  water  or  in  moist 
soil,  or  they  are  parasitic  on  plants  and  animals.^ 

^  [The  true  character  of  the  Arachnoidea  is  the  presence  of  four  pairs  of  ambulatory 
appendages.  This  number  is  reduced  to  two  pairs  in  the  gall-making  Phytoptid^^  and  they 
differ  from  all  other  Arthropoda  in  having  no  antennae. — F.  V.  T.] 

^  Twelve  orders  are  now  recognized,  as  follows  :  Pentastomida  or  Linguatulids  ;  Tardi- 
grada  or  bear-animalcules  ;  Phalangidce  or  harvest-men  ;  Acarina  or  ticks  and  mites  ; 
Palpigradi ;  Solifugce ;  Pseudoscorpiotiidea  or  book  mites;  Pedipalpi  or  false  scorpions; 
Scorpionidea  or  true  scorpions  ;  Aratieida  or  spiders ;  Xiphosura  or  king  crabs ;  and  Pycnogonida^ 
marine  Arachnoids. 

'  Chelifer  cancroides  has  also  been  observed  as  a  pseudoparasite  in  man  (Arnault  de 
Very,  S.,  "Pseudopar.  du.  Chel,  cancr.  chez  I'homme,"  Conipt.  rend.  Soc.  de  Biol.,  1901, 
liii,  p.  105). 

*  [The  chelicerae  are  sometimes  regarded  as  modified  antennae,  but  it  is  more  natural  to 
regard  them  as  the  morphological  equivalent  of  the  mandibles  of  Hexapoda. — F.  V.  T.] 

*  [The  pedipalpi,  or  second  pair  of  jaws,  consist  of  a  stout  basal  segment  and  a  palp,  which 
may  have  the  appearance  of  a  leg  in  Arachnida ;  this  may  end  with  or  without  a  claw,  or  with 
a  chela  (scorpions) ;  they  may  also  form  a  tube  enclosing  the  styliform  chelicerae  (mites). 
— F.  V.  T.] 

^  [Acarina  are  also  found  living  upon  trees,  feeding  upon  other  Arthropods  and  also  upon 
spores  of  lichen  and  fungi  {Oribatid(B  or  beetle  mites)  ;  they  also  swarm  indoors  amongst  stores 
and  provisions  {Tyroglyphidce  and  Glyciphagi,  household,  sugar  and  cheese  mites).  This 
order  is  very  important,  as  many  are  parasites  upon  man,  his  domestic  animals  and  his  cultivated 
plants,  and  attack  his  provisions  and  stores.  Some  live  on  blood,  and  in  some  of  the  ticks 
distribute  various  protozoal  and  other  blood  parasites  and  germs. — F.  V.  T.] 


TROMBIDIID^ 


48: 


Family.     Trombidiidae  (Running  Mites). 

Soft-skinned  Acarina  with  tracheae  and  with  two  eyes,  usually  pedunculated  ;  they 
are  often  brightly  coloured  ;  chelicer^e  lancet-  or  claw-shaped  ;  pedipalpi  claw-like  ; 
legs  composed  of  six  segments,  with  suctorial  discs  between  the  terminal  ungues.^ 
Larvae  six-legged.  To  the  latter  belong  the  larvae  of  several  species  of  Trombidium 
such  as  : — 

Genus.     Tronnbidium,  Latreille  (and  Leptus). 

Leptus  autumnalis,  Shaw,  1790. 

Leptus  occur  as  parasites  in  the  human  skin  and  cause  a  cutaneous 
disease  known  as  autumn  erythema,  and  produce  a  very  unpleasant 
sensation  on  account  of  the  trouble- 
some itching  ;  in  children  it  is  very 
often  accompanied  by  fever.^ 

Formerly  these  mites  were  considered 
adult  forms,  but  when  they  were  recog- 
nized as  mite  larvae  they  were  taken 
for  those  of  the  spider-mite  {Tetranychus 
telarius)  ;  the  investigations  of  Hanstein, 
however,    showed    this    to    be    a    mistake. 


Fig.   349. — Leptus  autumnalis,  with  so-called 
mcking    proboscis.     Enlarged.     (After  Gudden.) 


Fig.  350. — Leptus  autumnalis  :  the 
so-called  proboscis  is  formed  around 
the  hypopharynx  sunk  into  the  skin, 
loo/i.     (After  Trouessart.) 


When  Henking  first  investigated  the  development  of  Trombidium  fuliginosum, 
parasitic  in  the  larval  stage  on  vine-fretters,  he  demonstrated  the  occurrence  of  a 
form  very  similar  to  Leptus  autumnalis^  and  the  "autumn,  grass,  or  gooseberry" 
louse  was  commonly  designated  the  Trombidium  larva.  Even  before  Henking's 
work  it  had  been  described  by  Megnin  as  the  larva  of  Trombidium  holosericeum^  a 
red-coloured  species,  frequently  occurring  in  spring  and  summer  on  the  ground,  trees, 
etc.     This  assumption,  however,  as  Moniez  was  the  first  to  explain,  is  not  correct  ; 

^    [Some  have  seven  segments  to  the  legs. — F.  V.  T.] 

'  [This  minute  parasite  is  especially  obnoxious  in  barley  fields.  In  walking  over  barley 
stubble  one  is  sure  to  be  attacked  by  this  Acarus  in  many  districts.  Trombidium  is  often  very 
prevalent  in  gardens,  especially  along  rows  of  peas,  and  in  spring  they  may  be  found  on  fruit 
trees  and  bushes.  Nut-pickers  are  frequently  attacked  by  Leptus,  and  also  pickers  in  other 
fruit  plantations.     It  is  often  called  the  harvest  mite. — F.  V.  T.] 


486  THE   ANIMAL  PARASITES   OF   MAN 

indeed,  as  many  as  three  species  come  under  consideration  :  T.  gymnopterosum^ 
L.,  T.  fultginosufu,  Herm.  (according  to  Brucker),  and  two  species  known  hitherto 
only  in  the  early  stage,  T.  siriaticeps.  Helm,  et  Oudem.,  and  T.  poriceps,  Helm,  et 
Oudem.,  which  are  not  only  parasitic  on  mammals,  but  on  birds,  on  Arthropods  and 
especially  on  insects.     Arthropods  appear  to  be  the  normal  hosts  for  the  larvae. 

The  above-mentioned  forms  invade  the  skin  of  man  by  means 
of  their  oral  apparatus,  by  preference  invading  the  orifices  of  the 
sebaceous  glands  so  as  to  suck  the  blood  ;  around  the  point  attacked 
there  arises  a  wheal  about  the  size  of  a  lentil,  and  around  the  inserted 
hypopharynx  a  fibrinous  secretion,  the  ^'proboscis,"  which,  however, 
is  a  product  of  the  host,  just  as  chitinous  secretions  are  provoked  by 
Trombidia  parasitic  on  Arthropods. 

Further  species,  analogous  in  habit  to  Leptus  auiumnalis,  are  described  by  Riley 
from  Central  and  South  America  as  L.  americanus  and  L.  irritans. 

[L.  autumnalis  attacks  small  mammals  by  preference,  such  as  moles 
and  hares,  which  are  often  literally  covered  with  them.  Dogs  are 
also  subject  to  their  attack,  and  cats  suffer  similarly.  This  mite  also 
frequently  appears  in  colonies  on  cows ;  cavalry  horses  after  autumn 
manoeuvres  often  suffer  from  an  erythematous  affection  about  the 
hocks  and  knees  due  to  this  pest. 

[A  number  of  Leptus,  so  far  undescribed,  occur  abroad  which 
attack  man  in  the  same  way  as  L.  mitiimnalis  in  Europe.  Dr.  Durham 
has  brought  me  specimens  from  British  Guiana  called  hetc  rouge ; 
this  species  works  under  the  skin  much  as  does  our  European  species, 
but  it  is  very  distinct,  being  considerably  larger. — .p.  V.  T.] 


Trombldium  tialsahuate,  Lemaire,  1867. 

T.  tialsahuate  occurs  in  Mexico  under  conditions  similar  to  those 
of  Leptus  here.  It  also  frequently  attacks  men,  and  especially  fastens 
itself  on  to  the  eyelids,  in  the  axillae,  navel,  or  on  the  prepuce  ;  it 
induces  itching  and  swelling  of  the  parts  affected,  and  sometimes 
even  causes  suppuration ;  the  symptoms,  however,  generally  disappear 
after  a  week  and  remain  localized. ^ 


Other  species  of  mites  which  attack  man  are  reported,  mostly 
by  travellers,  from  various  other  places  ;  zoologically,  however, 
there  is  little  known  about  them.  The  pou  d'agouti  in  Guiana, 
niaibi  in  New  Granada,  colorada  in  Cuba,  mouqui  in  Para,  and  the 
buschmucker  in  New  Guinea  represent  a  few  of  these. 


'    Lemaire,  •'Import,  en  France  du  tialsahuate,"  Compt.  tend.  Acad.  Sci.,  Paris,   1867, 
Ixv,  p.  215. 


AKAMUSHI   OR   KEDANI 


487 


Akamushi  or  Kedani. 

In  a  few  districts  of  Japan  there  occurs  a  serious  illness,  with 
a  mortality  of  40  to  70  per  cent.  It  is  called  river  or  flood 
fever,  and  the  Japanese  doctors  have  connected  it  with  a  small 
mite  (akamushi,  kedani).  Baelz  has  opposed  this  opinion  on  the 
grounds  that  he  has  repeatedly  observed  the  same  species  of  mite 
in  his  dwelling  without  any  subsequent  illness  occurring.  Accord- 
ing to  Keisuke  Tanaka,  however,  a  connection  certainly  does  exist, 
inasmuch  as  the  akamushi,  like  Leptus,  attacks  persons  to  suck 
blood.  If  the  mite  is  not  removed,  or  if  the  spot  attacked  is 
injured    by   scratching,    etc.,    a   papule    surrounded   by   a   red    area 


Fig.  351. — The  kedani  mite.     Enlarged.     (After  Tanaka.) 

forms,  and  a  pustule  ensues  ;  and  finally  a  black  scab  covers  the 
seat  of  injury.  The  lesion  becomes  the  point  of  entrance  of  bacteria, 
especially  a  species  of  proteus  which  produces  river  fever.  If  the 
mites  are  carefully  removed  no  general  illness  takes  place. 

The  orange-red  mites,  which  we  only  know  in  their  larval 
condition,  measure  o*i6  to  0*58  mm.  in  length  by  o*io  to  0*24  mm. 
in  breadth.  They  have  leg-like  palpi  with  three  joints,  hirsute 
bodies,  and  very  hairy  legs  composed  of  five  segments,  terminating 
with  three  ungues. 


488 


THE   ANIMAL   PARASITES   OF   MAN 


Family.     Tetranychidae  (Spinning   Mites). 

These  have  tracheae  and  eyes  ;  the  palpi  are  composed  of  four  segments,  of 
which  the  last  but  one  has  a  powerful  claw.  The  legs  have  six  segments  with  sucker 
discs  between  the  claws. 

[Tiie  red  spiders  or  spinning  mites  (Tetranychi)  are  usually  placed 
in  the  family  Trombidiidce. — F.  V.  T.] 


Genus.     Tetranychus,  Dufour. 

Tetranychus  molestlssimus,  Weyenbergh,  1886.^ 

Found  in  Argentine  and  Uruguay  on  the  under  surface  of  the 
leaves  of  Xanthium  macrocarpiun ;  it  attacks  mammals  and  men, 
producing  severe  itching,  accompanied  by  fever  in  the  latter. 

It   has  been  asserted   by  Haller 
that  the  Cape  ailment  (Port  Natal 
;,    \v  M       sickness)  is  caused  by  mites,  but  this 

statement  has  been  contested. 

Tetranychus  telarius,  L.,  1758,^ 

var.  russeolus,  Koch. 

This  common  spinning  mite  like- 
wise attacks  human  beings,  but  the 
papules  produced  by  it  very  soon 
disappear. 

Family.     Tarsonemidae. 

A  family  distinguished  by  complete 
sexual  dimorphism,  the  species  of  which  are 
provided  with  tracheae ;  the  legs  have  five 
segments ;  the  terminal  segments  of  the 
front  pair  of  legs  of  both  sexes  possess  a 
claw  ;  the  terminal  segment  of  the  posterior 
pair  of  legs  of  the  male  likewise  has  a  claw. 
In  the   female  this   pair  of  legs,   like  the 

second  and  third  pairs  of  both  sexes,  is  provided  with  two  booklets  and  a  sucking 

disc.     The  cuticle  of  the  body  on  the  back  is  "  annulated." 

[This  family  of  small  transparent  mites  live  normally  as  plant 
parasites.  The  last  two  pairs  of  legs  are  widely  separated  from  the 
two  front  pairs. — F.  V.  T.] 

*  [This  species  is  also  known  as  Bicho  Colorado.  It  spins  a  web  under  the  lower  surface 
of  the  leaves,  and  it  is  only  from  December  to  February  that  it  attacks  warm-blooded  animals 
and  man. — F.  V.  T.] 

2  [There  is  something  wrong  here,  probably  in  the  identification.  T.  telarius  is  purely 
a  plant-feeder,  and  it  is  extremely  unlikely  a  variety  would  attack  man.  Anyhow,  it  will  not 
do  so  in  Great  Britain.— F.  V.  T.] 


Fig.   352.  —  7 eiranychus  telarius  var.  rus 
seolust  Koch.     Enlarged.     (After  Artault. 


PEDICULOIDES 


489 


Genus.     Pediculoides. 

Pediculoides  ventrlcosus,  Newport,  1850. 

Syn.:  Heteropus  ventrlcosus^  Newport,  1850;  Acarus  iritici^  Lagreze-Fossot, 
185 1  ;  Physogaster  larvarmn,  Lichtenstein,  1868;  Sphcerogyna  ventricosa,  Laboul- 
bene  and  Megnin,  1885. 

Males  are  oval  in  shape,  0*12  mm.  in  length  and  q-qS  mm.  in 
breadth,  flattened.  There  are  six  pairs  of  chitinous  hairs  on  the 
dorsal  surface  and  a  lyre-shaped  lamella  on  the  posterior  part.  The 
female  in  the  non-gravid  state  is  cylindrical  in  form,  0*2  mm.  in 
length  and  0-07  mm.  in  breadth  ;  when  gravid  the  posterior  part  of 
the  body  becomes  enlarged  into  a  ball,  which  may  attain  1-5  mm.  in 
size,  as  in  the  case  of  Pnlex  penetrans  and  of  the  female  Termites. 
On  emerging  the  young  are  already  provided  with  four  pairs  of  legs 
and  copulate  soon  after  birth. 


Fig.  353. — Pediculoides  ventricosus,     a,  male  ;  b,  young  female ;  c,  gravid  female. 
Enlarged.     (After  Laboulbene  and  Megnin.) 

These  animals  live  on  the  stalks  of  cereals,  and  feed  on  vegetable  and  animal 
juices  ;  they  are  also  found  on  corn-infesting  insects.  They  invade  the  barns  and 
seek  out  the  insects  living  in  the  dry  grains  of  corn,  or  wait  for  an  opportunity  of 
obtaining  food.  They  have  been  repeatedly  observed  on  human  beings,  particularly 
labourers  occupied  in  handling  grain  ;  their  bite  causes  severe  irritation,  local 
elevation  and  reddening  of  the  epidermis,  as  well  as  fever.  It  cannot  be  positively 
asserted  that  all  cases  of  the  occurrence  of  cereal  mites  on  man  relate  to  P.  ventrlcosus^ 
as  the  descriptions  are  often  insufficient.  Geber  states  that  one  form  is  Chrlthoptes 
7nonungulculosus^  or  Acarus  hordel;  Flemming  mentions  Tarsone?nus  unclnatus ; 
Koller  Orlbates  sp. ;  and  Karpelles  Tarsonemus  Intectus. 

[The  pregnant  female  Pediculoides  has  a  large  round  inflated  abdomen,  in 
which  the  ova  hatch  and  the  young  mature.  Later  they  escape  from  the  parent  as 
adults— F.  V.  T.] 


490 


THE   ANIMAL   PARASITES   OF   MAN 


Genus.     Nephrophages. 

Nephrophages  sanguinarius,  Miyake  and  Scriba,  1893. 

Males  measure  01 17  mm.  in  length  and  0-079  mm.  in  breadth; 
females  up  to  0*360  mm.  in  length  by  O'  1 20  mm.  in  breadth.  The  head 
is  provided  with  two  very  large  scissors-like  jaws  and  two  large  round 
eyes.  The  legs  are  composed  of  five  segments  and  are  all  of  equal 
length  ;  the  three  anterior  pairs  of  legs  have  pedunculated  ambulacra, 
the  posterior  ones  terminate  in  a  claw.  The  cuticle  on  the  back  is 
thickened  in  three  places,  shield-like;  the  abdominal  surface  without 
scutellum  is  longitudinally  striped  and  is  beset  with  chitinous  hairs. 

Colour  greenish  to  brownish- 


^^ 


s^ 


\ 


jr  '1        $*.    .       V  \ 


yellow.        Eggs 
0*040  mm. 


0*046     to 


Fig.  354. — Nephrophages  sanguinarius  :  male, 
ventral  surface.  Enlarged.  (After  Miyake  and 
Scriba.) 


Fig.  355. — Nephrophages  sanguin- 
arius :  female,  dorsal  aspect.  En- 
larged.    (After  Miyake  and  Scriba.) 


The  authors  discovered  these  mites,  but  always  dead,  in  the  urine  of  a  Japanese 
suffering  from  fibrinuria  complicated  with  chyluria  and  haematuria.  They  surmised 
that  they  were  endoparasites,  probably  situated  in  the  kidney  ;  but  this  view  is  not 
convincing,  though  they  also  report  that  for  a  week,  day  after  day,  the  mites  were 
found  in  the  patient's  urine,  as  well  as  in  urine  drawn  off  by  means  of  a  catheter, 
and  in  the  water  used  to  wash  out  the  bladder  (one  or  two  specimens  and  an  egg). 
The  statement  that  these  mites  have  large  eyes  makes  the  discovery  suspicious,  to 
say  the  least.  The  significance  of  the  discovery  is  not  supported  by  the  further 
statement  that  Disse  is  supposed  to  have  found  an  encapsuled  mite  closely  related 
to  the  Tyroglyphides  on  the  wall  of  the  vena  cava. 

In  the  case  of  Marpmann,  who  found  a  dead  Acarid  in  the  urine  of  a  man 
suffering  from  chronic  nephritis,  and  in  whom  later  examinations  proved  negative. 


EUPODID^,    GAMASID^ 


491 


the  author  himself  was  of  opinion  that  the  mite  had  reached  the  urine  from 
outside. 

We  are  certainly  acquainted  with  mites  living  endoparasitically,  namely,  the 
Cysticolce,  AnalgesincB^  of  which  Laminosioptes  galli?iarum  live  in  the  intramuscular 
and  subcutaneous  connective  tissue  of  fowls,  and  Cytoleichus  sarcopto'ides  in  their 
air  sacs.  Another  kind  of  mite  {Halarachne  halichoeri)  is  occasionally  found  in  the 
nasal  mucous  membrane  of  the  seal  {Haltchcerus  grypus)^  and,  quite  recently, 
Pneumonyssus  simicola^  which  is  more  nearly  related  to  Halarachne,  has  been  found 
in  the  lung  of  Cynocephalus  sp.  It  is  therefore  not  improbable  that  endoparasitic 
mites  are  found  in  man  ;  but  no  definite  discovery  has  yet  been  made. 

Family.     Eupodidae. 

Small  tracheate  mites,  with  moderately  long  or  short  pedipalpi,  composed  of 
four  segments,  of  which  the  last  segments  bend  ;  chelicer^e  forceps-shaped,  with 
serrated  edges  ;  legs  with  two  claws,  more  rarely  with  one,  and  terminating  in 
a  tuft  ornamented  with  fine  hairs;  genital  orifices  on  the  abdomen,  surrounded 
by  a  circle  of  little  hairs.  Most  species  live  free — one  lives  parasitically  on  the 
bodies  of  slugs. 

Genus.     Tydeus,    Koch. 

Tydeus  molestus,  Moniez,  1889. 

Male,  o'2  mm.  in  length,  o'i25  mm.  in  breadth.  Females,  0*225  mm. 
in  length,  o'i35  mm.  in  breadth;  gravid  female  0*315  to  0*360  mm. 
in  length  and  0*180  mm. 
in  breadth.  They  were 
observed  by  Moniez  on 
an  estate  in  Belgium, 
whither  the  creature 
had  apparently  been  im- 
ported twenty-five  years 
previously  with  Peru- 
vian guano  ;  they  ap- 
peared regularly  in  the 
summer  and  remained 
until  the  first  frost   set 

in;  they  were  found  on  grass-plots,  on  trees  and  bushes  in  masses; 
they  regularly  attacked  human  beings,  mammals  and  birds,  tormenting 
their  hosts  in  a  terrible  manner. 


Fig.  356.- 


Tydeus  molesHis  :    seen  in  profile. 
(After  Moniez.) 


Enlarged. 


Family.     Gamasidae  (Coleopterous  or   Insect  Mites). 

Chelicer^E  chelate  or  styliform  ;  pedipalpi  filiform  ;  the  legs  are  composed  of  six 
segments  with  two  terminal  ungues  and  a  bladder-like  sucking  disc  [caruncle — 
F.  V.  T.].  Stigmata  situated  between  the  third  and  fourth  pairs  of  legs  ;  the 
cuticle  thickened,  leather-like  ;  no  eyes  ;  the  larvas  have  six  legs. 

The  GamasidcB  are  predaceous  on  small  insects  and  other  mites  ;  some  are 
parasitic  on  insects,  and  one  is  noticeable  as  a  pest  on  birds,  etc. 

31 


492 


THE   ANIMAL   PARASITES   OF   MAN 


Genus.     Dermanyssus,  Dnges. 
Dermanyssus  gallinae,  de  Geer,  1778. 

Syn. :  Pnlex  gallincu,  Redi,  1674;  Atarus  gallincB^  de  Geer,  1778  ; 
Dermanyssus  avium^  Duges,  1834. 

The  male  measures  o*6  mm.  in  length  by  0-32  mm.  in  breadth;  the 
female  07  to  075  mm.  in  length  by  0*4  mm.  in  breadth.  The  body 
is  somewhat  pear-shaped  ;  the  colour  whitish,  reddish,  or  reddish- 
black,  according  to  the  contents  of  the  intestine.  The  legs  are. 
fairly  short  and  strong.  During  the  day  they  live  concealed  in  the 
nests,  cracks,  etc.,  of  the  hen-house,  and  at  night  attack  the  inmates 
in  order  to  suck  their  blood  ;  they  rarely  remain  long  on  the  birds. 
They  have  been  repeatedly  found  on  persons,  on  whose  skin  they 
produce  an  itching  eruption. 


Dermanyssus  hirundinis,   Hermann,   1804. 
Syn. :  Acarus  hirimdi?tts,  Herm.,  1804. 

Of  a  brownish  colour,  i-2  or  1*4  mm.  in  length  ;  lives  in  the  nests 
of  swallows  and  is  occasionally  found  on  man. 

[The  red  hen  mite  {Dermanyssus  galHnce)  not  only  attacks 
poultry  and   man,  as   stated  above,  but  is    found  on  all    birds  and 


Fig.  357. — Dermanyssus gallince.    En- 
larged.    (After  Berlese.) 


Fig.    358.  —  Dermanyssus     hirundinis. 
40/1.     (After  Delafond.) 


many  mammals.  The  D.  gallmce  is  the  same  as  D.  avium.  The 
species  found  in  swallows'  nests  is  also  said  to  be  the  same.  This 
mite  can  remain  for  weeks  without  any  food  from  its  normal 
host.  They  only  attack  man  when  entering  or  cleaning  dirty  and 
neglected  fowl-houses  ;  they  do  not  produce  a  true  dermatosis.  They, 
chiefly  attack  the  backs  of  the  hands  and  forearms  of  those  who  con- 
stantly attend  poultry  and  give  rise  to  symptoms  similar  to  the 
papular  eczema  of  scabies.     That  they  may  remain  some  time  upon 


IXODID^: 


493 


the  human  body  we  know  from  the  following  cases  out  of  many 
recorded :  Geber  observed  that  the  Dermanyssus  had  caused  a  diffused 
eczema  on  a  woman,  which  lasted  four  weeks  and  then  disappeared. 
The  tlque  of  F.  V.  Raspail  is  the  bird  Dermanyssus  ;  he  records 
children  and  adults  being  attacked  not  only  when  handling  pigeons, 
but  even  when  walking  in  a  garden  manured  with  pigeons'  dung. 
The  affection  soon  disappeared  wiien  the  pigeons  were  destroyed  and 
the  excreta  buried.  I  have  frequently  heard  of  poultrymen  being 
seriously  attacked  by  this  pest. — F.  V.  T.] 

Genus.     Holothyrus. 
Holothyrus  coccinella,  Gervais,  1842. 

Measures  5  mm.  in  size  ;  lives  on  birds  in  the  Island  of  Mauritius ; 
ducks  and  geese  frequently  fall  victims  to  its  bite  ;  it  also  attacks 
human  beings,  on  whose  skin  it  causes  severe  burning  and  swelling, 
but  no  reddening  ;  it  may  be  dangerous  to  children,  especially  by 
settling  in  the  oral  cavity. 

Other  Gamasides  occasionally  occur  in  man,  for  instance,  according  to  Moniez, 
Leignathiis  sylviariivi^  Canestr.  et  Fanzago  ;  according  to  Neumann  Lcelaps  siabularis. 
The  former  live  normally  in  the  nests  of  various  species  of  Sylvia^  Lselaps  on  dried 
vegetable  substances,  also  in  houses. 

[Marchoux  and  Conoy  {Bull.  Soc.  Path,  exot.,  1912,  v,  No.  10, 
pp.  796-798)  found  Leishman  granules  in  Lcelaps  echidnintis.  It  is 
assumed  that  Leishman  granules  may  be  found  in  most  Arachnoids, 
and  have  no  connection  wath  Spirochaeta. — F.  V.  T.] 

Family.     Ixodidae  (Ticks). 

Comparatively  large  Acarines  with  a  leathery  skin  ;  they  are  flattened  in  form, 
but  after  sucking  blood  the  abdomen  becomes  spherical  ;  the  cheliceras  are  rod-like 
and  possess  a  serrated  terminal  joint,  bent  hook-like  ;  the  median  parts  of  the 
pedipalpi  (maxillae)  form  a  rostrum  furnished  with  barbed  hooks  (fig.  359)  ;  the 
maxillary  palpi  themselves  are  club-like  or  rounded  ;  the  legs  are  composed  of  six 
segments  with  two  terminal  ungues,  often  also  with  "sucking  discs"  ;  the  stigmata 
are  at  the  sides  of  the  body,  posterior  to  the  fourth  or  third  pair  of  legs.  The  larvae 
are  six-legged. 

[The  true  ticks  (Ixodidcv)  are  all  blood-suckers,  and  as  far  as  is 
known  they  do  not  take  vegetable  food  at  all.  Not  only  are  the 
Ixodidcv  important  as  actual  parasites,  but  they  are  most  so  on 
account  of  the  fact  that  they  are  the  active  agents  in  carrying  various 
diseases  in  animals  and  apparently  in  man.  It  has  been  conclu- 
sively proved  that  the  bont  tick  {Amblyouiina  hebrceiun)  is  the  carrier 
of  the  fatal  ''heart-water  fever"  so  rife  amongst  sheep  in  South 
Africa,  that  the  dog  lick  {Hcvmaphy salts  leachi)  is  the  agent  by  which  the 
protozoa  that  cause  malignant  jaundice  in  dogs  are  distributed,  that 


494  THE   ANIMAL   PARASITES   OF   MAN 

Texas  fever  in  cattle  is  spread  by  Rhipicephahis  anniilattis,  and  Coast 
or  Rhodesian  fever  by  R.  appendiciilaUis  and  R.  siinns.  Their  im- 
portance as  disease  carriers  amongst  mammals  is  therefore  consider- 
able, and  it  may  prove  to  be  so  for  man.^  They  frequently  attack 
man,  but  chiefly,  according  to  my  observations,  in  their  early  stages 
in  Europe  ;  this  is  not  so,  however,  abroad.  The  life-history  of 
a  number  of  ticks  has  been  clearly  demonstrated.  Mr.  Wheler  has 
shown  that  in  Ixodes  reduvins  it  is  as  follows  :  the  female  deposits 
her  eggs  in  masses  upon  the  ground,  gradually  reducing  in  size 
as  the  eggs  pass  out,  until  she  finally  remains  a  mere  shrivelled 
empty  bag  and  then  dies.  The  eggs  are  oval,  golden  brown  in 
colour  and  smooth;  in  length  they  are  0*59  mm.;  as  in  all  Ixodidcv 
they  are  covered  with  a  glutinous  secretion,  by  means  of  w^hich 
they  adhere  together  in  masses.  These  egg  masses  may  be  deposited 
anywhere  on  the  ground,  but  amongst  rough,  coarse  herbage  seems 
to  be  the  favourite  place.  The  egg  stage  may  last  as  long  as 
twenty-two  weeks,  or  it  may  only  take  eight  weeks.  In  the  case 
of  the  bont  tick  a  single  female  may  deposit  15,000  or  more  eggs. 
The  process  of  egg-laying  is  described  as  follows  by  Mr.  Wheler  : 
"When  egg-laying  is  about  to  take  place,  the  head  is  further 
depressed  till  it  rests  close  against  the  under  side  of  the  body. 
In  this  attitude  the  end  of  the  rostrum  actually  touches  the  genital 
orifice,  the  palpi  being  at  the  same  time  widely  opened  out.  Behind 
the  head  and  from  beneath  the  shield,  at  what  for  the  purposes 
of  explanation  may  be  described  as  the  back  of  the  neck,  a  white, 
perfectly  transparent,  delicate  gelatinous  membrane  is  brought  dow^i 
through  inflation,  either  with  air  or  with  a  transparent  fluid,  above 
the  head,  which  it  temporarily  conceals.  The  end  of  this  mem- 
brane terminates  in  two  conical  points  which  appear  to  be  covered 
with  a  glutinous  secretion,  and  at  the  same  time  an  ovipositor  of  a 
somewhat  similar  character,  but  only  semi-transparent,  is  pushed 
forward  from  the  genital  orifice.  This  latter  is  a  tube,  within  which 
is  the  Qgg.  As  the  ovipositor  projects  it  turns  itself  inside  out,  like 
the  finger  of  a  glove,  leaving  the  egg  protruded  at  the  end  and  lying 
between  the  two  finger-like  points  of  the  membrane.  The  membrane 
and  the  ovipositor  are  then  withdrawn  each  from  the  other.  The 
tgg  adheres  to  the  former,  which  collapses  through  the  withdrawal 
of  its  contents,  dragging  the  head  forward  and  depositing  it  on  the 
top  of  the  head.  Neither  legs,  palpi,  nor  the  organs  of  the  mouth 
take  any  part  in  oviposition,  but  after  the  collapse  of  the  membrane 
the  palpi  are  closed  and  the  head  is  raised,  by  which  action  the 
egg  is  pushed  forward    to    the   front   edge    of    the    shield,    forming 

'  This  has  been  proved  in  Uganda— so-called  tick  fever  in  man. 


IXODID.E  495 

in  time  an  adherent  mass  of  eggs,  which  are  deposited  in  front  of 
the  tick." 

[The  egg  gives  rise  to  the  larval  form,  the  so-called  "seed-tick" 
stage.  At  first  these  minute  specks  are  pallid  and  soft,  but  they 
soon  harden  and  darken  in  colour.  These  larvae  are  six-legged  and 
crawl  up  grasses  and  various  plants,  and  there  await  a  passing  host, 
weaving  their  two  front  legs  in  the  air  and  becoming  attached  by  this 
means.  The  larval  ticks  feed  upon  the  blood  of  the  host,  and  when 
replete  fall  to  the  ground,  the  body  becoming  inflated  in  the 
meanwhile.  These  larvae  may  remain  on  the  host  only  two  days,  or 
they  may  remain  much  longer.  Eventually  they  moult  on  the  ground 
and  change  to  the  nymph  or  pupal  stage,  wiiich  has  four  pairs  of  legs. 
This  pupa  acts  just  as  the  larva,  crawls  up  plants  and  waits  to  regain 
the  host.  After  a  time  the  nymphs,  having  gorged  themselves  with 
blood,  fall  off  and  remain  on  the  ground  for  nearly  three  months  ; 
they  then  moult  and  become  adult  males  and  females.  In  about  ten 
days  they  assume  their  normal  colour  and  regain  the  host  afresh  ; 
the  female  gradually  swells  until  she  attains  that  large  inflated  form 
so  characteristic  of  ticks.  The  male  does  not  swell,  but  nevertheless 
feeds  upon  the  host  and  fertilizes  the  female. 

[The  act  of  coitus  is  strange :  the  male  tick  inserts  its  rostrum  and 
other  mouth  organs  into  the  sexual  orifice  of  the  female,  between  the 
base  of  the  posterior  pair  of  legs.  The  males  then  die  and  the 
females  fall  to  the  ground  and  deposit  the  ova.  There  are  variations 
in  the  different  species,  of  course,  from  those  given  above,  which 
apply  solely  to  Ixodes  reduvius.  The  larvae  and  nymphs  seem  to 
attack  most  animals,  but  the  adults  mainly  keep  to  the  same  host. 
The  periods  in  the  life-cycle  of  ticks  not  only  vary  in  the  different 
species,  but  in  each  species  according  to  climatic  conditions ;  for 
instance,  in  the  bont  tick  {Amblyounna  hehrceum,  Koch),  Lounsbury 
has  shown  that  the  development  is  rapid  in  summer,  slow  in  winter. 
The  period  from  the  time  that  the  female  drops  to  the  time  she 
commences  to  lay  eggs  varied  in  specimens  observed  by  him  from 
twelve  days  in  summer  to  twelve  weeks  in  winter,  and  the  complete 
period  from  the  dropping  of  the  female  to  the  hatching  of  the  eggs, 
from  eleven  weeks  in  summer  to  thirty-six  weeks  through  the  winter. 
Other  stages  vary  in  a  similar  manner. 

[Ticks  may  live  a  long  time  away  from  the  host  provided  they  are 
supplied  with  a  certain  amount  of  moisture.  Mr.  Wheler  kept  dog 
ticks  {Ixodes  pUunhetis)  in  the  larval  stage  for  ten  months  ;  the  pupae, 
male  and  female,  of  /.  reduvius  for  six  months. 

[I  have  kept  Oriiithodoms  mouhata  alive  for  eighteen  months 
without  food. 

[In  many  species  moulting  takes  place  off  the  host,  but  in  /.  hovis^ 


496  THE  ANIMAL   PARASITES   OF   MAN 

now  known  as  Rhipicephahis  annulatus^  Say  (the  carrier  of  Texas 
fever),  moulting  takes  place  on  the  host,  and  in  many  other  species 
also.^  Some  species  of  ticks  leave  their  host  on  its  death  (as  the  dog 
tick,  Hcemaphysalis  leaclii),  but  others  die  with  the  host  (bont  tick, 
Aniblyonifna  hebrceum). 

[Two  species  are  of  special  importance,  namely  Oniithodorus 
moubata,  Murray,  which  may  infect  human  beings  with  the  spirillum 
of  African  tick  fever,  and  Dermatocentor  reticulatus  var.  occidentalism 
which  is  said  to  be  the  carrier  of  Rocky  Mountain  spotted  fever. 

Classification  of  Ixodida\ 

[The  TICKS,  or  Ixodidce,  are  divided  into  two  groups,  known  as 
(1)  Argantiiicv,  (2)  Ixodince.  The  Argantince  are  told  from  the 
Ixodiiice  by  the  absence  of  dorsal  or  ventral  shields  in  both  sexes, 
and  also  by  the  rostrum  being  placed  beneath  the  cephalothorax, 
which  covers  it  over :  except  in  the  larval  stage,  in  which  it  is  sub- 
terminal,  and  in  the  pupal,  when  it  partly  projects.  Legs  nearly 
equal  in  length.  The  sexual  orifice  is  situated  between  the  two  first 
pairs  of  legs.     The  males  usually  smaller  than  the  females. 

[The  Ixodince  have  the  legs  unequal,  of  six  segments  with  two 
false  segments,  making  them  look  as  if  composed  of  eight  segments. 
The  rostrum  is  terminal  and  never  hidden  beneath  the  body. 
The  sexual  orifice  is  situated  between  the  bases  of  the  first  three 
pairs  of  legs.  In  the  males  the  orifice  is  obsolete  or  very  rudi- 
mentary, sexual  intercourse  being  effected  by  the  rostrum.  The 
males  are  smaller  than  the  females.  The  shield  in  the  females  never 
covers  so  much  as  one-half  of  the  body  even  when  fasting,  also 
in  the  larvae  and  nymphs ;  but  in  the  males,  which  do  not  distend, 
the  shield  covers  the  body  entirely,  or  all  but  a  narrow  margin. 
The  Ixodince  are  divided  into  two  groups  :  (i)  the  Ixodce,  and  (ii)  the 
Rhipicephalce.  The  former  have  a  long  proboscis  reaching  nearly  to 
the  end  of  the  palpi  or  even  a  little  longer  than  the  palpi.  The 
palpi  are  longer  than  broad.  The  Rhipicephalcv  have  short  palpi, 
nearly  or  quite  as  broad  as  long,  more  or  less  conical  or  subtriangular. 
They  were  called  Conipalpi  by  Canestrini. 

Synopsis  of  Genera. 

[A.  ArgantincE :  Rostrum  concealed  in  adult,  partly  exposed  in 
larvae  and  nymphs.     No  dorsal  and  ventral  shields. 
Body  flat  with    thin   edges,  finely  shagreened  and   punctate     Argas. 
Body  with  numerous    small  round  granules  and    with  thick 

sides  ...         ...         ...         ...         ...         ...         ...         ...     Ornithodorus. 

^  Some  ticks  require  only  one  [R.  decoloratits),  others  two  (A',  evertsi),  and  some 
three  hosts  {R.  appendiciilatus)  in  order  to  reach  maturity. 


IXODES 


497 


Body  with  dorsal  shield  over 


[B.  TxodmcE :  Rostrum  terminal, 
some  part  of  it. 

I.  Rostrum  and  palpi  longer  than  broad  (/;ir<?rt'^). 
o.  A  groove  around  anus  in  front. 

Palpi  caniculated  in  both  sexes      Ixodes. 

Palpiclaviform,  not  caniculated  in  the  male;  legs  very 

long Eschatocephalus. 

Palpi  claviform,  not  caniculated  in  the  male  ;  anal  groove 

absent  m  the  female  ...         ...         ...         ...         ...     Ceratixodes, 

fi.  A  groove  around  the  anus  behind. 

No  eyes ;  adanal  shields       ...         ...         ...         ...         ...     Apotiomma. 

Eyes  present. 

Males  with  no  adanal  shields  ...         ...         ...         ...     Atnblyoinma. 

Males  with  adanal  shields     ...         ...         ...         ...         ...     Hyalomma. 

II.  Labium  and  palpi  short;  palpi  not,  or  very  little,  longer 

than  broad  {^Rhipicephalce). 
o.  No  eyes. 

Rostrum  rectangular  ;  no  ventral  shields  in  the  male     ...     Hcemaphy salts, 
fi.  Eyes  present. 

No  adanal  shields,  but  usually  with  greatly  developed 
coxae  on  fourth  pair  of  legs.    Capitulum  quadrangular     Der7nacentor. 

Capitulum  hexagonal  ...         ...         ...         ...         ...     Rhipicentor. 

Adanal  shields  in  male.     Stigmata  comma-shaped        ...     Rhipicephalus. 

Stigmata  oval  or  round  ;  legs  normal        ...         ..;         ...     Boophilus. 

Segments  of  legs  expanded  Margaropus. 

[The  genus  Ceratixodes  of  Neumann,  1902,  occurs  on  birds. 
[The  genus  Eschatocephalus  of  Frauenfeld,  1853,  of  which  seven  species  are 
known,  is  mostly  parasitic  on  bats,  and  is  found  in  holes,  caves,  and  church  towers. 

[The    genus   Aponomma   of    Neumann,   1899,    is  exotic,   and  almost  entirely 
confined  to  snakes  and  saurians. 

[The  following  are  synonyms  of  the  different  genera : — 
Argas,    Latreille,  1796  {Rhynchoprz'on,  Hermann,  1804). 
Ixodes,   Latreille,    1795    {Acarus,    Linnaeus,    1758 ;     Cynorhcestes,    Hermann, 

1804  ;  Crotomts,  Dumeril,  1822). 
Ceratixodes,  Neumann,  1902  {Ixodes,  Cambridge,  1879  ;  Hyalom7na,  Cambridge, 

1879). 
Eschatocephalus,  Frauenfeld,  i^^2>{S'^^^onyssus,  Kolenati,  1857). 
Amblyomina,  Koch,  1844  {Ixodes,  Latreille,  1795). 
Hcemaphy sails,    Koch,    1844   {Rhipistoina,    Koch,    1844 ;    Go?itxodes,  Duges, 

1888  ;  Opitodon,  Canestrini,  1897). 
Rhipicephalus,  Koch,  1S44  {A carus,  Linnaeus,  1758;   Ixodes,  Latreille,  1795; 

Phanloixodes,  Berlese,  1889  ;  Boophilus,  Curtice,  1890). 
Dermacentor,  Koch,  1844  {Ixodes,  Latreille,  1795  5  Pseudixodes,  Haller,  1882). — 
F.  V.  T.] 

Genus.     Ixodes,  Latreille. 
Ixodes  reduvius,  L.,  1758.^ 

Syn. :  Acariis  reduvius  and  ricinus,  L.  ;  Ixodes  ricinus,  Latreille,  1806. 
The  males  are  oval ;  their  length  i*2  to  2  mm. ;  they  are  brownish- 
red  or  black  in  colour  ;  the  females  are  yellowish-red,  4  mm.  long; 

'  Ixodes  reduvius  and  /.  ricinus  are  synonymous.     [The  above  should  read  Ixodes  ricinus, 
Latreille,  1806.— F.  V.  T.] 


498 


THE   ANIMAL   PARASITES   OF   MAN 


when   gorged   they  are   lead-coloured,    and    may   attain    12    mm.  in 
length  by  6  to  7  mm.  in  breadth. 

The  dog  tick  (fig.  360)  lives  in  thickets  on  leaves,  etc.,  and  attacks  sheep  and 
oxen,  and  more  rarely  dogs,  horses,  and  human  beings,  into  the  skin  of  which  the 
female  bores  with  the  rostrum  in  order  to  suck  blood  ;  the  bite  is  not  dangerous,  and 
sometimes  is  not  even  felt.  Inflammation,  however,  is  set  up  if  the  creatures  are 
forcibly  removed  from  the  wound,  as  the  rostrum  as  a  rule  is  torn  off  in  the  process. 
If  left  alone  or  smeared  over  with  some  grease— vaseline,  oil,  butter,  etc. — the 
creatures  drop  off  spontaneously.  Sometimes  the  entire  tick  bores  itself  into  the 
skin  ;  they  also  appear  to  be  permanent  inmates  of  kennels. 

[The  species  /.  rednvius  is  the  same  as  /.  riciniis,  Latreille. 
The  male  is  2*35  to  2*80  mm.  long;  the  body  is  dark  brown,  almost 
black;  with  a  pale,  almost  white,  margin ;  there  are  also  traces 
of  reddish  mottling.  Coxae  of  the  first  pair  of  legs  with  a  short 
spine.  Rostrum  much  shorter  than  that  of  the  female  ;  shield  oval ; 
anal  shield  small,  about  one-third  the  length  of  ventral  shield.     The 

adult    female    varies    from   2*80 
to  3*5  mm.  when  not  distended, 
but    w^hen    gorged    may    reach 
A.^  )  ](    \  10  mm.  long.     The  shield  and 


Fig.  359. — A.,  the  rostrum  of  Ixodes 
ricinus  (male)  ;  B.,  the  terminal  joint  of  the 
maxillary  palpi  of  the  female.  Enlarged. 
(After  Pagenstecher.) 


Fig.  360. — Female  of  Ixodes  ricinus , 
gorged  full,  dorsal  and  ventral  surfaces. 
2/1.     (After  Pagenstecher.) 


legs  are  dark  blackish-brown,  body  deep  orange-red  with  four  dark 
longitudinal  lines,  paler  beneath  and  light  grey  in  front.  When  dis- 
tending it  is  pale  red  to  grey  or  white ;  when  fully  gorged  olive-green, 
or  dark  red  to  black,  with  irregular  yellow  streaks  on  the  back  and  sides 
just  before  egg-laying.  Sexual  orifice  opposite  fourth  pair  of  legs. 
The  nymph  varies  from  1*60  to  170  mm.  long  when  fasting;  the 
body  is  olive-white,  opaque,  with  four  distinct  brown  posterior 
markings  and  similar  anterior  ones,  leaving  a  pale  centre  to  the 
shield.  When  fully  gorged  it  is  3  mm.  long.  As  the  nymph  distends, 
it  changes  from  opaque  white  to  blue-black,  and  finally  black.  The 
little  larva  is  o'8o  to  1*50  mm.  long,  transparent  with  olive-green 
intestinal  markings ;  as  it  becomes  inflated  it  changes  to  blue-black, 
and  then  black.  There  are  no  eyes.  It  is  widely  distributed,  and 
chiefly  attacks  sheep ;  sometimes  it  occurs  on  dogs  and  also  attacks 


IXODES   HOLOCYCLUS  499 

man.  Megnin  records  it  from  horses  in  the  nymph  stage.  Amongst 
its  other  num.erous  hosts  are  goats,  cattle,  deer,  hedgehogs,  moles,  bats, 
birds,  and  lizards.  It  is  usually  known  as  the  grass  tick  and  bottle- 
nosed  tick.  This  species  occurs  in  Europe,  Asia,  North  Africa,  and 
North  America. 

[Synonyms. — Considerable  confusion  exists  over  the  name  of  this 
and  other  common  ticks,  owing  to  the  same  species  having  been 
described  under  a  great  many  names.  Observers  have  taken  the 
same  species  on  different  animals  and  in  various  stages  to  be  distinct, 
and  have  described  them  accordingly. 

[The  name  Ixodes  rednvitis,  Leach,  does  not  stand,  as  Leach  was 
describing  quite  a  different  parasite.  The  name  /.  ricinus,  Latreille, 
1806,  is  now  substituted  by  Neumann  and  Wheler. 

[The  synonyms  given  by  Wheler  are  as  follow  :  Rednvnis, 
Charleton,  1668 ;  Ricinns  caninus,  Ray,  1710  ;  Acarns  ricinoides, 
de  Geer,  1778  ;  Acarus  ricinns,  Linnaeus,  1788  ;  Cynorhcestes  rediiviiiSy 
Hermann,  1804;  Cynorhcestes  ricinns,  Hermann,  1804;  Ixodes 
inegatJiyrens,  Leach,  1815  ;  Ixodes  hipnnctatns,  Risso,  1826  ;  Cynorhcestes 
hermanni,  Risso,  1826;  Crotonns  ricinns,  Dumeril,  1829;  Ixodes 
traheatns,  Audouin,  1832  ;  Ixodes  plnmbens,  Duges,  1834  ;  Ixodes 
rednvitis,  Hahn,  1834;  Ixodes  fnscns,  Koch,  1835  (^)  J  Ixodes  lacertce, 
Koch,  1835  (?)  J  Ixodes pnstidarnni,  Lucas,  1866  ;  Ixodes  fodiens,  Murray, 
1877;  Ixodes  rnfus,  Ixodes  snlcatns,  and  Ixodes  scinri,  Koch. — F.  V.  T.] 

Ixodes  holocyclus,  Neumann,  1899. 

[Under  the  name  /.  holocyclns,  Cleland  {Jonrn.  Trop.  Med.  and  Hyg., 
1913,  xvi,  No.  3,  pp.  43-45)  says  that:  **  This  tick  is  common  in  man 
where  there  is  dense  scrub  and  tropical  jungle  along  the  east  coast  of 
Australia  at  certain  times  of  the  year.  It  may  cause  severe  symptoms 
in  children  resulting  in  death."  He  records  a  child  being  attacked  in 
1884  which  died,  and  another  case  from  which  200  ticks  were 
removed,  the  symptoms  being  weak  heart,  collapse,  syncope,  but  the 
patient  recovered  under  treatment ;  again,  in  the  same  journal 
(pp.  188,  189),  the  case  of  a  4i-year-old  girl  who  was  bitten  showed 
widespread  muscular  paralysis,  and  other  cases  resembling  conium 
poison. 

[Taylor  {Rep.  Ent.  Anst.  Inst.  Trop.  Med.,  191 1,  p.  21,  1913) 
refers  to  this  species  as  the  scrub  tick  of  New  South  Wales. 
The  partially  fed  female  has  a  dark  reddish-yellow  scutum  and  is 
almost  as  broad  as  long,  punctations  very  numerous,  equal  and 
confluent  in  places,  long  white  hairs  on  the  lower  half  of  each  coxa. 
He  records  it  as  attacking  man  commonly,  mentioning  Kamerunga, 
Cairns  district,  Queensland,  and  Sydney,  N.S.W.,  as  localities. — 
F.  V.  T.] 


500  THE    ANIMAL   PARASITES   OF   MAN 

Ixodes  hexagonus,  Leach,  1815. 
Syn.  :  Ixodes  sexpunctatus,  Koch,  1897;  /.  viilpis,  Pagenstecher,  1861. 

Lives  in  the  same  manner  as  the  foregoing;  especially  attacks 
hounds,  but  also  other  mammals  and  even  birds.  The  difference 
consists  in  the  shape  of  the  legs,  the  shorter  rostrum,  and  the  larger 
size  of  the  male.  It  also  occasionally  attacks  man,  but  is  usually 
confused  with  the  previously  mentioned  species. 

[The  synonyms  of  this  species  are  as  follow  :  Ixodes  antiimnaliSj 
Leach,  1815;  /.  m//(^c^/,  Audouin,  1832;  /.  rediivhis,  Audouin,  1832; 
/.  crennlatns,  Koch;  /.  erinaceus,  Murray,  1877;  /.  ricinus,  Megnin, 
1880.     Two  other  synonyms  are  given  above  by  Braun. 

[The  female  when  fully  replete  is  11  mm.  long,  when  fasting 
3-86  mm. ;  the  shield  is  heart-shaped  and  punctate,  body  finely  hairy  ; 
palpi  short  and  broad ;  labium  shorter,  and  tarsi  of  all  the  legs  more 
truncate  than  in  /.  ricinus.  The  colour  of  the  distended  body  is 
drab  and  somewhat  waxy ;  rostrum,  shield  and  legs  light  testaceous. 
The  male  varies  from  3*5  to  4*0  mm.  long,  and  is  reddish-brown 
in  colour  with  lighter  legs;  the  shield  is  punctate  and  leaves  a 
narrow  margin  around  the  body  ;  the  body  is  elliptical,  almost  as 
large  in  front  as  behind.  There  is  a  spine  on  the  coxae  of  the  first 
pair  of  legs,  which  is  shorter  than  in  the  m.ale  /.  ricinus  and  longer 
than  in  the  female.  The  genital  orifice  is  opposite  the  interval 
between  the  second  and  third  pair  of  legs.  The  fasting  nymph  is 
176  mm.  long,  light  bluish-grey,  margined  and  transparent,  with 
four  large  posterior  intestinal  marks  joined  together  behind  the 
shield  and  smaller  ones  extending  to  the  front  and  sides.  When 
fully  distended  it  is  uniformly  brownish-white ;  shield,  legs  and 
rostrum  pale  testaceous.  The  larva  varies  from  o"88  mm.  when 
fasting  to  176  mm.  when  gorged.  Its  body  is  light,  but  gradually 
becomes  darker,  with  similar  intestinal  marks  to  ricinus. 

[This  tick  is  very  common,  especially  on  ferrets,  stoats  and  hedge- 
hogs. It  is  also  found  on  sheep,  cattle,  etc.  The  males  do  not 
generally  occur  in  company  with  the  females  on  the  host.  Pairing 
probably  takes  place  on  the  ground. — F.  V.  T.] 

Genus.     Amblyomma,  Koch. 

Amblyomma  cayennense,  Koch,  1844. 

Syn.  :  Amblyomma  mixtum^  Koch,  1844  ;  Ixodes  herrerce^  Duges,  1887  ; 
Amblyomma  scuiptum^  Berlese,  1888. 

Characterized  by  the  possession  of  eyes.  The  male  measures 
3-8  mm.  in  length  by  3  mm.  in  breadth ;  the  female  4  mm.  in  length 
by  3  mm.  in  breadth,  but  when   full  of  blood  may  become  13  mm. 

^  Neumann,  G.  L.,  "Rev.  de  la  fam.  des  Ixodides,"  ///  Mem.  Soc.  Zool.  France,  1899, 
xii,    p.  129. 


HYALOMMA  501 

in  length  and  11  mm.  in  breadth.  They  are  common  in  the  whole 
of  Central  America  (Carrapatas);  and  attack  mammals,  amphibious 
animals  and  man.^ 

[This  species  was  described  by  Fabricius.  It  occurs  in  Cayenne, 
Guiana,  in  Southern  Texas,  Florida,  California,  Mexico,  Guatemala, 
Honduras,  Nicaragua,  Costa  Rica,  Panama,  Bermuda,  Cuba,  Jamaica, 
Trinidad,  Colombia,  Venezuela,  French  Guiana,  Brazil,  Paraguay 
and  the  Argentine.  It  is  called  the  silver  tick.  It  frequently  attacks 
man.  Schwarz  and  Bishopp  {Bull.  105,  U.S.  Dept.  Agric,  p.  158) 
heard  of  one  man  w^hose  legs  were  well  covered  with  suppurating 
sores  and  w^ho  was  ill  from  the  attack  of  these  ticks  and  the  wounds 
produced  by  scratching,  and  records  other  cases  of  their  swarming 
on  man.  Newstead  {Ann.  T.rop.  Med.  and  Par.,  1909,  iii,  No.  4,  p.  442) 
records  it  as  the  worst  pest  to  man  in  Jamaica. — F.  V.  T.] 

Amblyomma  americana,    Linnaeus. 

The  so-called  long  star  tick,  from  the  silvery  spot  on  the  apex  of 
the  scutum  of  the  female.  It  will  attack  any  mammal  and  even  birds 
and  also  man.  It  occurs  in  North  America,  and  also  in  Brazil, 
Guiana  and  Guatemala.  Its  punctures  frequently  end  in  suppuration. 
In  the  Eastern  and  Southern  States  man  is  more  frequently  attacked 
by  this  species  than  any  other.  Moss-gatherers  in  Louisiana  are 
badly  attacked  by  it.^  It  also  attacks  the  milkers  in  dairies.  Attempts 
to  transmit  Texas  fever  failed  with  this  species. 

Amblyomma  maculatum,  Koch. 

The  so-called  Gulf  Coast  tick,  of  the  Gulf  Coast,  occurs  on  birds,^ 
mammals  and  man,  especially  cattle,  and  attacks  the  ears. 

Genus.     Hyalomma,  Koch. 

Hyalomma  aegyptium,  L.,  1758. 
Syn.  :  Acariis  cegyptius^  L.,  1758  ;  Ixodes  camelinus^  Fischer,  1823. 
A  species  frequently  found  in  Africa,  particularly  in  Egypt  and 
Algeria,  and  which  also  occurs  in  France  and  Italy,  as  well  as  in 
Asia.  Male  8  mm.  in  length,  4*5  mm.  in  breadth.  Female  up  ta 
24  mm.  in  length  and  15  mm.  in  breadth.  It  infests  large  and  small 
animals  as  well  as  human  beings.^ 

[This  is  one  of  the  largest  ticks,  nearly  reaching  the  size  of  the 
bont  tick.     It  is  known  in  Africa  as  the  bont  leg-tick;  all  farm  stock 

^  Neumann,  G.  Y^.^Joc.  cit.^  p.  205. 

2  Morgan,  "  Ticks  and  Texas  Fever,"  Louisiana  State  Bull.  55,  pp.  134,  135,  P^-  59- 

3  Neumann,  G.  L.,  loc.  cit.,  p.  285  ;  Ronsisvalle,  "  Sui  fenomeni  morb.  prodotti  nel  uomo 
da  un  Ixodide  denominate  Hyal.  ceg.,''  Boll.  Ace.  Gioenia  sci.  vat.,  1891,  xvii. 


502  THE   ANIMAL   PARASITES   OF   MAN 

is  attacked,  but  sheep  and  goats  suffer  most.  Only  one  generation 
appears  to  occur  each  year.  The  male  is  almost  black  with  a  pale 
marginal  stripe;  the  replete  female  brown  with  irregular  light  blue 
stripes.     It  is  abundant  in  parts  of  South  Africa.] 

Genus.     Haemaphysalls,  Koch. 
Haemaphysalis  punctata,  Canestrini  and  Fanzago,  1877 — 1878. 

Syn.  :  Hcemaphysalis  sulcata^  Canestrini  and  Fanzago,  1877 — 1878  ;  Rhico- 
cephaltis  exposiiicius,  Koch,  1877  ;  HcEinaphysalis  peregriniis^  Cambridge,  1889  ; 
Herpetobia  sulcata,  Canestrini,  1890. 

[This  species  does  not  appear  to  be  common.  It  occurs  on  sheep, 
goats,  horses  and  cattle.  I  have  seen  a  female  taken  from  man  in 
Britain.  The  female  when  fasting  is  3*44  mm.  long,  when  gorged 
12  mm.  long.  Colour,  reddish-brown,  leaden -grey  when  gorged; 
dorsal  shield  deeply  indented  in  front ;  rostrum,  shield  and  legs 
brownish ;  body  finely  punctate,  both  above  and  below ;  sexual 
opening  opposite  the  coxae  of  the  second  pair  of  legs  in  both  sexes. 
Palpi  a  little  longer  than  the  labium ;  first  segment  short  and  narrow, 
second  and  third  widened  on  the  dorsal  face.  Coxae  with  a  short, 
broad  blunt  spine  ;  tarsi  short,  terminated  with  a  spur  on  the  first 
pair.  The  male  is  3*10  mm.  long.  Body  rather  narrow,  yellowish 
to  reddish-brown  ;  dorsal  shield  nearly  covers  the  whole  body; 
numerous  punctures  over  the  whole  surface.  Eleven  indentations  on 
the  posterior  margin  of  the  body  ;  peritremes  lighter  in  colour,  large 
and  comma-shaped.  The  three  anterior  pairs  of  legs  with  a  short 
spine  on  the  haunches,  the  fourth  with  a  very  long  one  directed 
backwards.  The  nymph  varies  from  2*5  to  3*0  mm.,  is  oval,  and 
light  yellow  to  dark  red  in  colour.  Dorsal  shield  rounded  with  few 
punctations.  No  spur  on  tarsi,  and  sexual  orifice  nearly  obsolete. 
Larva  short  and  oval.     Length  1*20  mm. — F.  V.  T.] 

Genus.     Dermacentor,  Koch. 

Dermacentor  reticulatus,  Fabricius,  1794. 

Syn.  :  Acariis  reticulatus,  Fabr.,  1794;  Ixodes  reticulatus,  Latreille,  1806; 
/.  uiarmoratus,  Risso,  1826. 

This  tick  is  provided  with  eyes,  but  it  is  distinguished  from  Ixodes 
and  analogous  genera  by  the  lack  of  the  abdominal  plastron  in  the 
male,  which  measures  5  to  6  mm.  in  length  by  3-5  mm.  in  breadth. 
The  female  may  attain  16  mm.  in  length  and  10  mm.  in  breadth. 
It  is  found  in  the  South  of  Europe,  in  Asia,  and  in  America; 
it    attacks   chiefly  oxen,    sheep   and    goats,    and    occasionally  man.^ 

'  Neumann,  G.  L.,  "  Rev.  de  la  fam.  des  Ixodides,"  J//w.  Soc.  Zool.  France,  1897,  x,  p.  360. 


DERMACENTOR   VENUSTUS  503 

[This  tick  sometimes  causes  much  annoyance  to  human  beings. 
It  was  once  most  troublesome  at  Revelstoke.  Specimens  have 
recently  been  found  on  fowls,  turkeys  and  pheasants  in  Kent. 

[Other  synonyms  are  as  follow  :  Cynorhcestes  pichiSf  Hermanur 
1804;  Crotonus  variegatiis,  Dumeril,  1829;  I.  pictus,  Gervais,  1844; 
Deiinacentor  albicolUs,  Koch,  1844 — 1847;  D.  pardalinus,  Koch,  1844 
— 1847;  ^'  ferruglneus,  Koch,  1844 — 1847;  Ixodes  holsattis,  Kolenati, 
1857  >  Psetidixodes-holsatuSf  Haller,  1882  ;  Hcemaphysalis  inarmorata, 
Berlese,  1887. 

[The  female  when  fasting  is  3'86  mm.  long  by  2  mm.  wide. 
The  body  is  depressed,  larger  behind  and  reddish -brown  in  colour. 
The  shield  is  very  large  and  extends  to  the  level  of  the  third  pair 
of  legs,  with  a  few  large  and  many  small  punctations,  milky  white, 
variegated  with  reddish-brown.  Sexual  orifice  opposite  the  coxae 
of  the  second  pair  of  legs.  Coxae  of  the  front  legs  are  deeply 
bifid,  the  others  with  a  moderate  spine.  When  gorged  light 
brown,  and  may  reach  16  mm.  When  depositing  eggs  the  female 
is  mottled  with  dark  brow^n  above  and  below.  The  male  is  like 
the  female.  The  shield  is  reddish-brown,  variegated  with  a  milky 
white  pattern.  Coxae  of  the  fourth  pair  of  legs  three  times  the 
size  of  the  third.  There  is  a  sharp  backwardly  pointing  spine  on 
the  second  palpal  segment,  also  seen  (but  smaller)  in  the  female. 
Length  4*20  mm. 

[According  to  Mr.  Wheier  this  is  a  very  variable  species  both 
in  size  and  colour.  It  occurs  in  England  on  sheep,  but  not  com- 
monly. It  has  probably  been  introduced  into  Britain.  Besides 
the  animals  mentioned  above  it  is  also  found  on  deer. —  F.  V.  T.] 

Dermacentor  venustus,  Banks. 

[The  Rocky  Mountain  tick  fever  tick.  This  species  has  been 
wrongly  called  Dermatocentor  reticidaUis  var.  occidentalis.  The  correct 
name  of  the  carrier  of  Rocky  Mountain  tick  fever  is  Dermacentor 
venustus,  Banks  (Hooker,  Bishopp  and  Wood,  Bidl.  106,  U.S.  Dept. 
Agric,  Div.  Ent.,  p.  165). 

[The  female  is  from  i3'8  by  10  by  6*4  mm.  to  16*5  by  11*4  by 
6*9  mm.  when  gorged ;  the  male  from  2*1  by  1*5  mm.  to  6  by 
3*7  by  1*4  mm.  The  male  reddish-brown  ;  scutum  with  an  exten- 
sive pattern  of  white  lines,  usually  but  little  white  on  the  mid- 
posterior  region,  legs  slightly  lighter  than  scutum,  joints  tipped  with 
white.  Female  with  scutum  mostly  covered  with  white,  abdomen 
reddish-brown,  legs  as  in  male.  The  nymph  when  unengorged 
reddish-brown,  when  gorged  dark  bluish-grey ;  the  larva  is  yellowish- 
brown  when  unengorged,  slate  blue  when  engorged.  The  ova  light 
brown,  shinv  and  smooth. 


504 


THE   ANIMAL   PARASITES   OF   MAN 


[The  chief  wild  hosts  are  the  brown  bear,  coyote,  woodchuck, 
rabbit,  wild  cat,  badger  and  mountain  goat  for  the  larvae  ;  practically 
all  small  mammals  act  as  hosts  for  larvae  and  nymphae,  whilst  the 
adults  are  seldom  found  on  other  than  large  domestic  animals  ;  horses 
and  cattle  are  preferred.  It  occurs  in  British  Columbia,  southward 
to  Northern  New  Mexico,  and  from  the  foothills  of  the  Rocky 
Mountains  in  Colorado  to  the  base  of  the  Cascade  Range  in  Oregon 
and  California;  abundant  in  Western  Montana,  Idaho,  Eastern 
Washington,  Oregon,  North  Utah,  West  Wyoming  and  North-west 
Colorado. 

[Of  great  importance  in  the  Bitter  Root  Valley  of  Montana,  where 
a  number  of  cases  of  fever  occur  each  year,  with  a  mortality  of  about 
70  per  cent.  In  British  Columbia  this  tick  causes  tick  paralysis  in 
man  and  sheep.  Only  the  adults  seem  to  attack  man  and  animals 
there  (Hadwen  and  Nuttall,  Parasitology,  1913,  vi,  No.  3,  pp.  288- 
297  and  298-301)  according  to  the  Canadian  Medical  Associa- 
tion Jonrnal,  December,  1912.  The  symptoms  are  unlike  spotted 
fever.  For  full  details  of  this  tick  vide  Bulls.  105  and  106,  U.S. 
Dept.  Agric] 

Dermacentor  occidentalis,  Neumann. 

This  tick  only  occurs  in  the  Pacific  Coast  region  of  the  United 
States.  Owing  to  the  fact  that  it  frequently  attacks  man  as  well  as 
occurring  in  great  abundance  in  Oregon  and  California,  it  is  of  con- 
siderable economic  importance.  It  is  spoken  of  as  the  wood  tick,  and 
in  the  regions  where  found  is  the  most  common  tick  to  attack  man. 
Hooker,  Bishopp  and  Wood  {Bnll.  106,  U.S.  Dept.  Agric,  Div.  Ent., 
1912,  p.  189)  state  that  a  number  of  cases  have  been  brought  to  their 
notice  where  the  bite  of  this  tick  has  caused  considerable  local  inflam- 
mation, which  in  some  cases  required  physicians'  attention.  It  has 
been  supposed  to  be  connected  with  Rocky  Mountain  spotted  fever, 
but  it  is  doubtful  if  it  is  concerned  in  its  transmission.  The  engorged 
female  is  steel  grey,  the  dorsum  with  an  olive-green  surface  colour, 
which  covers  the  grey  except  in  small  spots,  giving  a  mottled 
appearance.  The  unengorged  males  and  females  are  reddish-brown, 
scutum  covered  with  a  whitish  bloom,  interrupted  by  many  red 
punctures.  The  female  is  9  by  6-i  by  3-3  mm.  to  ii'S  by  y6  by 
5-6  mm.  ;  the  male  2*8  by  i-6  mm.  to  4*2  by  2-3  mm.  The  larvre 
are  bluish-grey  when  engorged,  reddish-brown  when  unengorged. 
The  nymph  is  light  brown,  sides  of  scutum  darker,  and  the 
intestines  dark  brown.  It  is  confined  to  the  Coast  Range  and 
Sierra  Nevada  Mountains  in  California  and  Oregon  and  the  small 
mountain  range  to  the  south-west. 


MARGAROPUS,    RHIPICEPHALUS,    SPECIES   OF   ARGAS  505 

Dermacentor  variabilis,  Say. 
The  American  dog  tick  has  also  been  found  on  man,  but  it  is  of 
Httle  economic  importance  as  it  is  easily  removed  from  its  host. 

Genus.     Margaropus,  Karsch. 
Margaropus  annulatus  australis,  Fuller. 
The  so-called  Australian  cattle  tick.     Newstead  ^  reports  this  as  a 
great  pest  to  man  in  Jamaica  in  its  larval  stage.     Its  chief  hosts  are 
cattle,  horses,  goats,  sheep,  dogs  and  rabbits. 

Margaropus  microplus,  Canestrini. 
Recorded  by  Aragao  (Mem.  Inst.  Osivaldo  Cruz,  1911,  iii,  fasc.  2, 
p.  163)  as  occurring  in  larval  stage  on  man  in  Brazil. 

Genus.      Rhipicephalus,  Koch. 

Rhipicephalus  sanguineus,  Latreille,  1804. 

Syn. :  Ixodes  sanguineus,  Latr.,  1804;  /.  rufiis,  Koch,  1844  ;  Rhipicephalus  limbatus^ 

Koch,  1844  ;  Rh.  siculus,  Koch,  1844  ;  Rh.  stigmaticus,  Gerstacker,  1873. 

Spread  over  almost  the  entire  tropical  and  sub-tropical  regions, 

occurring  in  Europe  in  the  South  of  France  and  in  Italy  ;  it  infests 

dogs  and  more  rarely  sheep;  oxen,  cats,  foxes   and    human   beings 

are  also  attacked.^ 

Neumann's  Table  of  Species  of  Argas. 

J  Body  elliptical  (sides  curved)     2. 

i  Body  oblong  (sides  straight),  ending  anteriorly  in  a  point  ...  7. 

J  Body  transversely  oval vespertilionis. 

(  Body  elongate  oval  3. 

f  Margin  of  body  striated 4. 

t  Margin  of  body  formed  by  quadrangular  areolae       PERSICUS. 

{Body  flat,  integument  plainly  wrinkled  ...         ...         ...  5. 

Body  tumid,  elongate  ;  integument  finely  wrinkled  ;  coxae  of 

fourth  pair  of  legs  near  anterior  third  of  body        ...         ...  hennanii. 

J  Body  oval,  narrowed  anteriorly rEflexus. 

5    (  Body  elliptical,  blunt,  hardly  narrowed  anteriorly 6. 

f  Body  twice  as  long  as  broad      ...         ...         ...  cucu77ierinus. 

\  Body  hardly  longer  than  broad transgariepinus. 

Dorsal  integument  with  large  polygonal  depressions  ;  tarsi 

appearing  bifid brumptii. 

Dorsal   integument   almost   smooth  ;    tarsi    not    appearing 

bifid         ...         ...         ...         ...         ...         ...         ...         ...  cBqualis. 

The  Argantince  are  distinguished  from  the  Ixodince  by  the  head,  which  in 
the  former  is  situated  on  the  inferior  aspect  of  the  '  cephalothorax,  while 
in  the  Ixodince  it  projects  freely  ;  also  by  the  very  short  proboscis,  the  small 
club-like  palpi,  the  lack  of  suckers  on  the  legs,  as  well  as  by  the  scutellum,  which 
covers  the  entire  back  and  is  bent  up  round  the  borders.  Two  genera  are  dis- 
tinguished :  Argas,  Latreille,  1796  (Rhynchoprion,  Hermann,  1804),  and  Ornithodorus, 
Koch,  1844,     The  species  live  on  mammals,  but  more  especially  on  birds. 

'  Ann.   Trop.  Med.  and  Par.,  1909,  iii,  No.  4.      ^  Neumann,  G.  L.,  he.  cit.,  1897,  p.  385. 


5o6  THE   ANIMAL   PARASITES   OF  MAN 

Genus.     Argas,    Lalreille. 
Argas  reflexus,  Fabricius,  1794. 

Syn.  :  Acarus  7'cjlexus,  Fabricius,  1794  ;  A.  ina7'gi7iatus,  Fabricius,  1794; 
Rhy7ichoprio7i  coluDibcr^  Hermann,  1804. 

The  European  marginated  tick,   Argas   reflexus    (length  of   male 
4   mm.,   breadth   3   mm.,    length    of    female   6   to   8   mm.,   breadth 

4  mm.),  is  of  a  yellowish  colour  and  has  yellowish-white  legs.  The 
ingested  blood  shows  red  or  brown  through  the  intestine,  which  is 
provided  with  blind  sacs.  It  lives  in  dovecots.  It  remains  hidden 
during  the  day  and  at  night  crawls  on  to  the  sleeping  pigeons  to  suck 
their  blood.  It  has  been  observed  in  France,  England,  Italy,  Germany, 
and  Russia.  Persons  sleeping  near  infected  dovecots,  or  in  apart- 
ments formed  from  pigeon-lofts,  are  also  attacked,  even  when  the 
room  in  question  has  not  been  used  for  sheltering  pigeons  for  years, 
as  '^marginated  ticks"  can  live  in  a  fasting  condition  for  a  very  long 
time.  The  bite  sometimes  gives  rise  to  serious  symptoms,  such  as 
general  erythema  and  sudden  oedema. 

[This  pest  more  often  feeds  on  the  blood 
of  man  than  is  imagined.  Blanchard  states 
that  he  has  received  them  from  men's  clothes 
in  Strasburg.  Boschulte,  of  Westphalia,  re- 
cords these  parasites  in  a  bedroom  inhabited 
by  children  and  connected  with  a  pigeon- 
house.  The  children  were  bitten  during  sleep 
on  the  hands  and  feet.  The  result  of  the  bite 
was  intense  itching  along  the  nerves,  the  bite 
^      ^       ,  ^  only  beini^  marked  by  a  red  spot.     In  a  girl 

from  the  dorsal  surface,  the  of  14  or  15,  vesiclcs  wcrc  lormed  Similar  to 
intestine  showing  through     thosc  produced  by  burns,  and  in   an  old  man 

the     integuments.        (After  ^  n        ,^.,  1         •     r    1 

Pagenstecher.)  an  ulcer  formed.     Others  record  painful  punc- 

tures and  persistent  oedema  produced  by  this 
pigeon  pest.  It  was  once  abundant  in  Canterbury  Cathedral,  and 
often  caused  much  annoyance,  I  am  told,  to  the  worshippers  ;  the 
ticks  falling  down  from  the  roof,  where  they  were  living,  deriv^ed 
from  the  numerous  pigeons  that  breed  in  the  towers.  This  Acarus 
has  enormous  powers  of  vitality,  living  without  food  for  months  at 
a  time.— F.  V.  T.] 

Argas  persicus,  Fischer  de  Waldheim,  1824. 

Of  oval  form  and  brownish-red  colour.     The  male  measures  4  to 

5  mm.  in  length  by  3  mm.  in  breadth  ;  the  female  7  to  10  mm.  in 
length  by  5  to  6  mm.  in  breadth.     It  frequents  the  entire  north-west 


ARGAS   BRUMPTI  507 

and  north-east  of  Persia  (the  gerib-gez  or  malleh  of  the  Persians,  the 
miana  bug  of  travellers).  It  lives  concealed  in  houses  and  attacks 
man  at  night  to  suck  his  blood.  Its  bite  is  much  dreaded,  but  the 
serious  results  may  probably  be  attributed  to  unsuitable  treatment  of 
the  wound  or  its  invasion   by  bacteria. 

[This  tick,  sometimes  called  the  tampan  and  wandlius  in  South 
Africa,  is  mainly  a  fowl  parasite.  Fow4s  and  ducks  frequently  die 
under  its  attack,  particularly  young  ones,  death  being  due  to  loss 
of  blood.  This  tick  remains  attached  to  its  host  during  its  larval 
stage  for  about  five  days  ;  it  then  leaves  and  moults  in  concealment. 
In  its  subsequent  stages  it  visits  its  host  by  night  and  remains  for 
about  an  hour  only,  during  which  time  it  distends  itself  fully  with 
blood.  As  a  nymph  it  moults  twice,  not  once  as  do  the  cattle  ticks. 
This  tick  and  other  Argas  become  larger  with  each  moult,  but  retain 
their  same  general  appearance.  The  female  visits  the  host  every  now 
and  then,  and,  between,  deposits  eggs  in  sheltered  crevices.  About 
fifty  to  120  are  deposited  at  once.  Four  weeks  seems  a  necessary 
period  to  intervene  between  visits  to  the  host,  and  the  interval  may 
be  extended  to  upwards  of  a  year  according  to  Lounsbury.'* 

[It  is  found  in  the  Sudan,  where  Balfour 
has  found  granules  derived  from  the  seg- 
mentation of  spirilla  in  their  digestive  tract. 
Fantham  and  Hindi  have  confirmed  this.  It 
has  been  assumed  that  these  granules  carry 
infection. 

[This  so-called  Persian  tick,  the  miana, 
which  is  such  a  scourge  to  travellers  in  Persia, 
appears  to  infest  the  huts  of  natives  in  that 
country.  It  has  been  sent  me  from  Quetta, 
where  it  has  invaded  houses  to  such  an  extent       Fig.  362.— ^;x«.f  perskus: 

.  .  i  1-        •      ii  rrM  •      1  ventral   aspect.      7/1.     (After 

the  natives  cannot  live  in  them.    1  he  virulence     Megnin.) 
of  its  bite  is  probably  due  to  the  tick  trans- 
mitting fever  germs  from  natives,  probably  inured,  to  strangers,  who 
would  be  susceptible. — F.  V.  T.] 


Argas  brumpti,  Neumann. 

[Found  in  Somaliland,  by  Brumpt,  and  in  the  Sudan.  This  tick 
attacks  man  as  well  as  wild  animals  and  produces  a  painful  swelling, 
according  to  King,^  but  as  pointed  out  by  that  naturalist  it  probably 
relies  on  other  than  human  food. — F.  V.  T.] 


'   «*  Report  of  Government  Entomologist,  Cape  of  Good  Hope,  for  1899,"  1900,  p.  ZZ- 
2  "Fourth  Report  Wellcome  Res.  Labs.,"  1911,  p.  128. 
32 


5o8 


THE   ANIMAL   PARASITES   OF   MAN 


Argas  chinche,  Gervais,  1844. 

This   Acarus,  a  native    of   the   temperate   parts   of   Colombia,   is 
very  troublesome  to  man.    It  is  probably  identical  with  A.  americaiuiSf 

Packard,  which  infests  domestic  fowls  and  turkeys,  and  occasionally 
also  cattle,  and  is  differentiated  from  A.  reflexus  by  the  sculpturing 
of  the  cuticle. 

Genus.     Ornithodorus,  Koch. 

Neumann's    Synopsis    of    the    Genus    Ornithodorus    is    as 
follows  :  — 

Hypostome  unarmed ;  integument  in  nymph  stage  and  partly 

in  adult  spinulose ...  megnini. 

Hypostome    armed    with    recurved    teeth  ;    integument    not 

spinulose     ...         ...         ...         ...         ...         ...  2. 

Camerostome  with  movable  lateral  flaps talaje. 

Camerostome  without  movable  lateral  flaps       ...         ...         ...  3. 

■  Anterior  border  of  distal  segments  of  legs  with  tubercles  or 

festoons       4. 

Anterior  border   of  segments   of  legs   without    tubercles   or 

festoons       ...         ...         ...         ...         ...         ...         ...         ...  8. 

Body  not  much  contracted  anteriorly       5. 

Body  pointed  anteriorly       ...         ...         ...         ...         ...         ...  7. 

Tubercles  of  distal  segments  of  legs  higher  than  broad,  distant  6. 
Festoons  of  distal  segments  of  legs  as  broad  as  high,   con- 
tiguous        ...         ...         ...         ...         ...         ...         ...         ...  pavimefiiosus. 

Eyes  present SAVIGNYI. 

1  No  eyes           moubata, 

J  Eyes  present coriaceus. 

1  No  eyes          turicata. 

f  Integument  with  fine  radiating  wrinkles lahorensis, 

\  Integument  granular            9. 

j  Tarsi  appearing  bifid  at  apex         furcostts. 

1  Tarsi  not  appearing  bifid  at  apex 10. 

(  Tarsi  of  first  pair  of  legs  with  three  dorsal  tubercles,  of  other 

\       legs  with  one          caTiestrinii. 

(^  Tarsi  without  dorsal  tubercles  or  with  only  one            11. 

Tarsi   of  last   three   pairs   of  legs   with    pronounced    dorsal 

protuberance          tholozanii. 

.  Tarsi  of  legs  with  indistinct  dorsal  protuberance          errattcus. 


Ornithodorus  moubata,  Murray,  1877. 
An  abundant  African  tick  which  is  one  of  the  caixiers  of  the 
spirillum  of  African  relapsing  fever  and  can  also  carry  Filaria  perstans 
(Christy).  Its  body  is  oval,  yellowish-brown  when  young,  greenish- 
brown  when  mature  The  integument  is  covered  with  mamillose 
tubercles.  No  eyes  jid  the  stout  legs  granular  above,  the  tibiae  and 
tarsi  fringed  with  tubercles  on  the  upper  side.    Pocock  ^  records  it  from 


1  *•  A  System  of  Medicine,"  AUbutt  and  RoUeston,  i,  pt.  2,  p.  195. 


ORNITHODORUS   SAVIGNYI,  CORIACEUS,  TALAJE,  AND   TURICATA      509 

Uganda  and  German  East  Africa,  Congo  and  Angola,  to  Namaqualand 
and  the  Transv'aal  in  the  south.  It  is  called  bibo  in  Uganda,  moiibata 
in  Angola,  and  tampan  on  the  Lower  Zambesi.  It  feeds  on  animals 
and  birds  as  well  as  man.  Its  bite  is  very  painful.  This  tick  is  found 
in  native  huts,  living  in  cracks  and  crevices  and  in  the  thatch  roofs. 

The  female  tick  infected  with  the  spirillum  transmits  the  infection 
to  the  eggs  and  the  next  generation.  They  appear  to  be  able  to  live 
without  food  a  long  time,  and  probably  live  for  years.  They  lay  their 
eggs  in  masses  on  the  ground  or  in  crevices,  and  when  they  hatch 
they  are  in  the  nymph  stage  with  four  pairs  of  legs.  0.  moubata  also 
occurs  in  Madagascar  with  recurrent  fever  (Lamoureux,  Bull.  Soc. 
Path,  exot.,  1913,  vi,  No.  3,  pp.  146-149). 

Ornlthodorus  savignyi,  Audouin,  1827. 

At  one  time  considered  the  same  as  the  preceding  species,  but  can 
be  easily  separated  by  the  presence  of  two  pairs  of  eyes.  It  is  widely 
spread  over  Africa  and  has  been  found  in  South  India  and  at  Aden. 
In  the  Sudan  it  occurs  in  large  numbers.  King  ^  records  that  a  few 
miles  N.N.E.  of  Khartoum  370  specimens  were  collected  in  two 
hours  under  a  single  tree  by  a  well.  It  is  found  in  Somaliland, 
where  relapsing  fever  occurs  and  no  0.  moiibata^  which  it  probably 
replaces  as  a  transmitter  (Drake-Brockman,  "  Rep.  Col.  Office,"  April  6 
and  April  16,  1913).  It  also  occurs  in  Tunis,  where  the  natives  call  it 
"tobbiah"  (Weiss,  Arch,  de  I' Inst.  Pasteur  de  Tnnis,  1912,  pt.  4, 
p.  226). 

Ornlthodorus  coriaceus,  Koch. 
Found  in  Mexico,  Paraguay  and  California.     Attacks  man. 

Ornithodorus  talaje,  Guerin,  1849. 

An  eyeless  species  with  somewhat  elongate  pentagonal  body  found 
in  Mexico  and  South  America,  called  the  '^chinche."  A  variety  of 
it  {coniceps)  is  found  at  Venice,  etc.,  and  another  variety  on  various 
islands  in  the  Indian  Ocean  and  South  Atlantic.  Its  bite  is  very 
painful  to  man. 

Ornithodorus  turicata,  Duges,  1876. 

Without  eyes.  Indigenous  in  Central  America  ;  attacks  human 
beings  and  pigs.  The  bite  is  painful  and  is  often  followed  by 
serious  consequences. 

[So  virulent  is  this  species  that  pigs  put  in  an  infested  sty  often 
die  in  a  night.  This  "  turicatas "  of  Mexico  often  reaches  6  mm. 
in  length.— F.  V.  T.] 

'  "Fourth  Report  Wellcome  Res.  Labs.,"  191 1,  B,  p.  129. 


510 


THE   ANIMAL    PARASITES    OF   MAN 


Ornithodorus  tholozani,   Laboulbene  and  IVregnin,   1882. 
Syn. :  Argas  tholozani^  Lab.  and  Meg.,  1882. 

Without  eyes.  Males  4  to  6  mm.  in  length  and  2  to  4  mm.  in 
breadth  ;  females  8  to  10  mm.  in  length  and  4  to  5  mm.  in  breadth. 
It  especially  attacks  sheep.     Native  of  Persia  and  Asia  Minor. 

[This  species  is  reputed  as  being  very  dangerous  to  man.  It  is 
locally  know  as  the  kene,  or  sheep-bug.  In  its  fully  gorged  state  it  is 
deep  violet.— F.  V.  T.] 

Ornithodorus  megnini,  Duges,  1883. 

Syn.  :  Argas  viegnini^  Duges,  1883. 

Length  8*5  mm.,  breadth  5*5  mm.     Native  of  Mexico. 

[Another  synonym  for  this  species  is  Rhynclioprion  spinosiun, 
Marx.  The  adult  males  and  females  are  grey  to  dark  brown,  the 
male  somewhat  the  smaller  ;  female  5  by  3*5  by  2-5  mm.  to  10  by 
6  by  3*5  mm.  The  larvae  at  the  seed  tick  stage  are  dark  grey, 
turning  to  pink,  then  to  a  whitish  grey  when  engorged.  The  nymph 
when  young  is  blood-red  in  front,  rest  pearly  white  ;  later  they  turn 
reddish-brown. 

[Intense  pain  may  be  caused  by  its  presence  in  and  around 
the  ears. 

[Two  specimens  in  the  nymphal  state  were  taken  from  the  ears  of 
a  visitor  to  Cambridge  by  Dr.  ].  Christian  Simpson.  They  were 
supposed  to  have  entered  the  ears  when  the  gentleman  was  camping 
out  in  Arizona  (Lancet,  1901,  i,  No.  4,052,  p.  1198,  fig.  3). 

[This  species  attacks  the  horse,  ass,  dog,  cats  and  oxen,  generally 
around  the  ears,  and  also  attacks  man.  It  is  well  known  in  the 
United  States  as  infesting  the  ears  of  children  (New  York  Ent.  Soc. 
Journ.,  1893,  pp.  49-52). 

[It  occurs  in  Texas,  Arizona,  New  Mexico  and  California  as  well 
as  Mexico,  Brazil,  and  possibly  many  parts  of  South  America  ;  and 
recently  Bedford  (*'  Sec.  Report  Div.  Vet.  Res.,  S.  African  Union," 
1912,  pp.  343,  344)  has  shown  it  to  occur  at  Vryburg  and  Fauresmith, 
in  the  Transvaal,  on  stock.     It  also  occurs  in  the  Sudan. — F.  V.  T.] 

Other  Literature  on  Ixodida. 

(i)  "  Penetration  de  V Ixodes  ricinns  sous  la  peau  de  rhomme,"  Conipt.  rend.  Soc.  de  Biol.^ 
1891,  xliii,  ser.  9,  iii,  pp.  689-691,  R.  Blanchard. 

(2)  "Notas  sobre  Ixodidas  brazileiros, "  Mem.  Inst.  Oszvaldo  Cruz,  1911,  iii*  fasc.  2, 
pp.  145-195,  pis.  II  and  12,  Dr.  H.  de  Beaurepaire  Aragao.     Table  of  Brazilian  Species. 

(3)  **  Contribuicao  para  a  sistematica  e  biolojia  dos  Ixodidas,"  Alein.  Inst.  Oswaldo  Cruz., 
1912,  iv,  fasc.  I,  pp.  96-120,  pis.  2  and  3,  Dr.  H.  de  Beaurepaire  Aragao. 


TYROGLYPHID^  511 

Family.     Tyroglyphidae. 

Very  small  mites  without  eyes  and  without  tracheae,  with  smooth  skin. 

The  males  usually  have  a  suctorial  pore  on  either  side  of  the  anus,  which  is  used 
during  copulation,  or  suckers  may  be  found  in  both  sexes  near  the  sexual  orifice. 
The  mouth  parts  form  a  cone  with  chelate  cheliceras,  and  three-jointed  pedipalpi  ; 
the  legs  are  usually  short,  have  five  segments  with  a  terminal  claw  and  suckers,  or 
either  one  or  other  of  these  organs.  The  numerous  species  and  genera  live  free  and 
from  choice  in  slowly  decomposing  vegetable  and  animal  matter  (cheese,  cereals,  flour, 
sugar,  presel'ves,  dried  anatomical  preparations,  bacon,  dried  fruits  and  fungi),  also  in 
the  corners  of  dwellings,  elc.  ;  they  incidentally  get  into  or  on  to  man,  or  are  found  in 
chamber  utensils  and  in  spittoons  ;  actual  parasites  are  rarely  found  amongst  them. 

[The  chief  genera  are  Tyroglyphus,  Rhizoglyphus,  Glyciphagus,  Aleurobius 
and  Histiogaster.  The  first  three  have  typical  characters  referred  to,  but  are 
distinguished  from  each  other  by  the  two  former  having  the  hairs  on  the  dorsum 
smooth,  whilst  in  Glyciphagus  they  are  hairy,  plumose,  or  feathered.  Rhizoglyphus 
can  be  told  from  Tyroglyphus  by  having  claws  on  the  tarsi  without  any  suckers  ; 
Tyroglyphus  has  both  claws  and  suckers.  The  larvae  are  hexapod  and  may  become 
adult  in  the  usual  way  by  repeated  moults,  or  they  enter  the  so-called  hypopial 
stage.  In  this  the  eight-legged  nymph  becomes  quiescent,  and  during  this  stage 
it  fixes  itself  to  some  insect  or  other  animal  by  a  patch  of  suckers  on  the  lower 
surface  of  the  hind  end  of  the  body,  and  is  so  carried  from  place  to  place.  The 
hypopus  does  not  feed  and  has  a  hard  shell  and  short  legs.  When  it  has  reached 
a  new  home  it  moults  and  development  proceeds  in  the  normal  way.  Canestrini 
and  Kramer  treat  the  TyroglyphidcB  as  a  sub-family  of  the  Sarcoptidce^  calling  them 
sub-family  Tyroglyphi7Ke,i\\Q  other  sub-families  being  Sarcoptince,  CanestriniiticE  and 
Analsmce. — F.  V.  T.] 

Sub-family.     Tyroglyphinae. 

Genus.     Aleurobius,  Canestrini. 

Aleurobius  (Tyroglyphus)  farinae,  de  Geer  (part),  Koch. 

The  male  measures  o'33  mm.  in  length  by  o*i6  mm.  in  breadth; 
the  female  o'6  mm.  in  length  by  0*3  mm.  in  breadth.  These  mites 
possess  five  pairs  of  suctorial  organs  of  a  light  colour  ;  the  legs  are 
reddish.  Moniez  observed  them  in  Lille  on  the  skin  of  labourers 
who  had  been  unloading  Russian  corn.  A  few  of  the  species 
generally  mentioned  under  the  designation  of  Tyroglyphus  slro  are 
probably  the  common  flour-mite,  which  also  occurs  on  dry  cheese. 

[The  farince  of  de  Geer  is  an  Aleurobius  described  by  him  in  1778 
("  Mem.  Hist.  Ins.,"  vii,  t.  5,  f.  15,  p.  97)  as  Acariis  farincv. — F.  V.  T.] 

Genus.     Tyroglyphus,  Latreille. 

Tyroglyphus  siro,  L.,  1756. 

(Defined  by  Gervais,  1844.) 

Male  0-5  mm.  in  length  by  0*25  mm.  in  breadth  ;  female  0-53  mm. 

in  length  by  0*28  mm.  in  breadth;  the  males  have  two  suckers  on  the 

tarsi    of    the   fourth   pair  of  legs.     Penis  straight,  colour  whitish  or 

reddish. 


512 


THE   ANIMAL   PARASITES   OF   MAN 


Tyroglyphus  longior,  Gervais,  1844. 

White  or  yellowish,  with  two  black  spots  on  the  abdomen. 
Male  o*55  mm.  in  length,  0*28  mm.  in  breadth  ;  penis  bent. 
Female  o*6i  mm.  in  length  and  0*28  mm.  in  breadth. 

r.  siro  and  T.  longior  live  on  dry  cheese,  in  flour,  on  dried  fruits, 
etc.,  and  have  been  occasionally  observed  in  the  stools,  urine,  or  pus 
of  human  beings,  and  also  on  their  skin.  The  so-called  vanillismus 
is  to  be  attributed  to  these  species. 

[T.  siro  and  T.farincc  of  Schrank  (non  Geer)  are  the  same.  They 
are  described  under  other  names,  such  as  Acarus  lactis,  Linn. ; 
A.  favormrij  Herm.,  etc.  ;  A.  lactis  in  milk,  farince  in  flour,  and  siro 
in  cheese;  and  as  A.  dysenterice,  Linnaeus  (^' Syst.  Nat.,"  ed.  12, 
pp.  1024-1767).] 


Fig.  363. —  TyroglypJnis  fariuLC  :  male. 
Enlarged.     (After  Berlese.) 


Fig.     364.  —   Tyroglyphus'  longior^ 
Gerv.     (After  Fum.  and  Robin.) 


It  is  to  these  species  that  a  case  of  dysentery  was  referred. 
Rolander,  who  studied  under  Linnaeus,  was  attacked  by  what  was 
called  dysentery.  The  complaint  soon  gave  way  to  treatment, 
but  eight  days  after  it  returned,  soon  disappeared,  but  again 
came  a  third  time.  All  the  time  Rolander  had  been  living  like 
the  other  inmates  of  the  house,  who  all  escaped.  Linnaeus,  aware 
that  Bartholemy  had  attributed  dysentery  to  insects  which  he 
said  he  had  seen,  advised  his  student  to  examine  his  stool.  The 
result  was  that  innumerable  mites  were  found  to  be  present.  Their 
presence  was  easily  accounted  for  by  the  fact  that  they  were  found 
in  numbers  in  a  cup  made  of  juniper  wood  from  which  the  student 
alone  drank  of  a  night,  and  they  were  found  to  be  of  the  same  species. 
What  this  species  is  we  do  not  know.  Linnaeus  called  it  Acarus 
dyseniericv,  but  it  was  the  same  as  his  Acarus  siro.     No  records  have 


GLYCIPHAGUS  513 

occurred  since.  It  cannot  be,  as  Latreille  supposed,  the  cheese  mite, 
for  they  have  been  eaten  by  milhons  since,  and  it  is  strange  no  such 
case  has  occurred  again. 

[Tyroglyphus  minor  var.  Castellani,  Hirst, 

causes  the  copra  itch  in  persons  employed  in  the  copra  mills  in 
Ceylon.  The  skin  of  the  hands,  arms,  legs  and  even  body 
becomes  covered  with  pruriginous  papules,  papulo-pustules  and 
pustules  near  the  head.  The  eruption  begins  as  a  rule  on  the 
liands.  The  mites  live  in  the  copra  dust.  They  produce  dermatitis. 
Castellani  produced  the  disease  experimentally  by  rubbing  copra 
dust  containing  mites  on  the  skin  of  healthy  people.  Beta-naphthol 
ointment  (5  to  10  percent.)  proved  useful  in  treatment  (Jonrii.  Trop. 
Med.  and  Hyg.,  December  16,  191 2,  Castellani  and  Hirst). — F.  V.  T.] 

Genus.     Glyciphagus,  Hering,  1838. 
Glyciphagus  prunorum,  Her.,  and  G.  domesticus,  de  Geer. 

The  Glyciphagi  are  differentiated  from  the  Tyroglyphi  in  that 
the  chitinous  hairs  on  the  body  are  fringed  or  feathered,  and  that 
they  lack  a  furrow  dividing  the  cephalothorax  from  the  abdomen. 
They  live  under  similar  conditions  to  the  Tyroglyphi  and  are 
occasionally  found  on  man  or  in  faeces. 

[Sugar  merchants  and  grocers  are  frequently  troubled  by  swarms 
of  G.  dojuestiais,  which  leave  the  stores  when  being  handled,  and 
especially  shopmen,  who  handle  sugar  kept  in  small  stores  for  some 
time.  These  are  the  Acari  that  cause  that  irritating  temporary  affec- 
tion known  as  "  grocer's  itch." — F.  V.  T.] 

Glyciphagus  cursor,  Gervais. 

Under  this  name  Signor  Moriggia  figures  a  horny  excrescence 
of  great  length  growing  from  a  woman's  hand,  and  containing  in 
its  cavities  quantities  of  Acarus.  This  species  is  really  G.  domesticus, 
de  Geer.  G.  domesticus  has  also  been  described  by  Gervais  (Ann.  Sci. 
Nat.f  1841,  ser.  2,  xv,  p.  8)  as  G.  hippopodes. 

Glyciphagus  buski,  Murray.^ 

[This  is  a  mite  found  by  Busk  and  named  after  him  by  Murray. 
It  was  taken  from  beneath  the  cuticle  of  the  sole  of  the  foot  of 
a  negro  in  the  Seamen's  Hospital  Ship  on  the  Thames  in  1841, 
in  large  sores  of  a  peculiar  character  confined  to  the   soles  of  the 

'  Cooper  and  Busk's  Micros.  Jonrn.,  1842,  and  "Economic  Entomology,"  Murray, 
p.  280. 


5H 


THE   ANIMAL   PARASITES   OF  MAN 


feet.  It  appeared  that  the  disease  was  caused  by  its  burrowing 
beneath  the  thick  cuticle.  The  disease  was  attributed  to  the 
wearing  of  a  pair  of  shoes  which  had  been  lent  to  another  negro 
whose  feet  had  been  similarly  affected  for  nearly  a  year.  The  negro 
to  whom  the  shoes  were  lent  came  from  Sierra  Leone.  Mr.  Busk 
stated  that  some  water  brought  by  Dr.  Stranger  from  the  River 
Sinoe,  on  the  coast  of  Africa,  contained  one  nearly  perfect  specimen, 
and  fragments  of  others  very  similar  to  if  not  identical  with  this 
Acarus.  Mr.  Busk  adds  that  he  had  been  informed  by  Staff- 
Assistant  Surgeon  P.  D.  Murray  that  at  Sierra  Leone  there  is  a 
native  pustular  disease  called  craw-craw — a  species  of  itch  breaking 
into  open  sores. 

[From  Busk's  original  figure  I  see  no  reason  to  doubt  that  this 
is  a  Glyciphagus.— F.  V.  T.] 


Genus.     Rhizoglyphus,  Claparede,  iJ 
Rhizoglyphus  parasiticus,  Dalgetty,  1901. 

The  Rhizoglyphii  are  to  be  recognized  by  their  short  legs,  which  are  beset 
with  spines,  and  by  the  tarsi,  which  terminate  in  a  claw.  They  live  on  plants, 
roots  and  bulbs,  especially  the  bulbs  of  lilies. 


YiG.  T,es.— Rhizoglyphus parasitiats.     a,,  male  ;  ^.,  female.       Enlarged.   (After  Dalgetly.) 

This  species  has  been  observed  on  the  feet  of  Indian  coolies 
working  in  the  tea  plantations;  they  produce  a  skin  disease 
which  always  commences  with  blebs  between  the  toes,  and  which 
almost   always    extends    to    the    malleoli,     but     not    beyond.      The 


HISTIOGASTER 


5^5 


Acari  have  an  elliptical  body,  which  is  grey,  but  varies  from 
greenish-yellow  to  greenish-brown  when  the  stomach  is  full.  Eyes 
are  absent.  The  legs  are  composed  of  five  segments  and  terminate 
with  a  claw.  The  males  measure  o'i8  mm.  in  length  by  o'o8  mm. 
in  breadth,  and  possess  genital  and  anal  pores  ;  the  females  measure 
0'2  mm.  in  length  by  0*09  mm.  in  breadth.^  [This  is  also  known  as 
coolie  itch  and  is  common  in  Indian  tea  plantations. — F.  V.  T.] 

Genus.     Histiogaster,  Berlese,  1883. 
Histiogaster  (entomophagus?)  spermaticus,  Trouessart,  1900. 

The  genus  Histiogaster,  which  also  approaches  the  Tyroglyphince^  is 
characterized  by  the  circumstance  that  the  males  possess  suctorial  pores 
used  in  copulation,  as  well  as  leaf-shaped  appendages  at  the  posterior  end 
of  the  body.     They  feed  on  vegetables,  especially  on  small  fungi. 


Fig.  366. — Histiogaster  (entomophagus  ?)  spermaticus :  on  left,  male;  on  light,  female — both 
from  the  abdominal  aspect.     200/ 1.     (After  E.  Trouessart.) 

This  species  has  been  described  by  Trouessart,^  who  found 
numerous  specimens,  some  adult,  others  in  the  developmental  stage 
(larvae,  nymphs),   and  ova,  in  the  fluid  removed    by  puncture   from 

^  Dalgetty,  A.  B.,  "Water-itch;  or  Sore  Feet  of  Coolies,"  Journ.  Trop.  Med., 
1901,  iv,  p.  73. 

^  Trouessart,  E.  R.,  Comft.  renJ.  Soc.  Biol.,  Paris,  1900,  lii,  pp.  742-744,  893,  894; 
Arch,  de  Par.,  1902,  v,  pp.  449-459. 


5l6  THE   ANIMAL   PARASITES   OP^   MAN 

a  cyst  of  the  right  testis.  The  males  measure  0-25  mm.,  the  females 
0*32  mm.,  and  the  larvae  o'l  mm.  in  length.  The  author  is  of  opinion 
that  the  animal — perhaps  a  fertilized  female — was  introduced  by  a 
catheter,  and,  as  a  matter  of  fact,  it  was  afterwards  found  that  the 
patient  had  once  had  the  catheter  passed  in  India  while  suffering  from 
pernicious  fever. 

It  would  here  rather  appear  to  be  the  case  of  a  facultative  para- 
sitism of  an  otherwise  free-living  species.  Histiogastcr  ciitoiiiopliagtts, 
Laboulbene,  is  found  occasionally  in  collections  of  insects  feeding 
on  larger  species  containing  much  fat  ;  the  species  also  occurs  on  dry 
cantharides;  it  appears  to  belong  to  the  region  of  South  Europe, 
where,  however,  it  is  widely  spread. 

[Entomophagus  occurs  all  over  Europe  and  in  America.  It  has 
been  described  under  the  following  names  :  Acanis  tnaliis,  Shimer,  1868 
{Trans.  Illinois  Hort.  Soc.)  ;  Dernialeichus  niali,  Riley,  1873  {Rep.  Ins. 
Missonri,  v,  p.  87) ;  Tyroglyphns  mali,  Murray,  1877  ("  Eco.  Ent.  Apt.," 
p.  275)  ;  T.  coriicalis,  Michael,  1885  {Trans.  Roy.  Micros.  Soc,  ser.  2,  v, 
3,  p.  27,  figs.  I  to  14)  ;  Histiogaster  coriicalis,  Canestrini,  1888 
{Prosp.  Acarof.,  iii,  p.  397)  ;  H.  alenrophagns,  Sicherin,  1894,  Canestrini, 
Prosp.  Acarof.,  vi,  p.  815.  Trouessart's  species  is  evidently  distinct. 
— F.  V.  T.] 

Genus.     Cheyletus. 

Cheyletus  nnericourti,  Lab. 

Acaropsis  mericourti^  Moq.  Tand. 
[This  mite  has  been  described  from  three  specimens  found  in  pus 
which  flowed  from  an  abscess  in  the  ear  of  a  naval  officer,  produced 
by  inflammation  of  the  auditory  passage.  Where  the  mites  came 
from  we  do  not  know,  as  they  were  found  near  the  Bank  of  New- 
foundland. This  genus  of  Acari  has  enormous  mandibles  and  a 
peculiar  tracheal  system  ;  two  ungues  and  appendages  to  the  tarsi. — 
F.  V.  T.] 

Family.     Sarcoptidae  (Itch  Mites). 

Small  mites  without  eyes  and  tracheae,  and  with  delicate,  transversely  striated 
cuticle.  The  mouth  parts  form  a  cone,  over  which  the  shield-shaped  upper  lip 
protrudes  ;  the  chelicerae  are  chelate  ;  the  pedipalpi  (or  maxillary  palpi)  have  three 
joints  ;  the  legs  are  short  and  compact,  and  composed  of  five  segments  ;  the  terminal 
joints  have  pedunculated  suckers  (ambulacra)  or  a  long  bristle.  The  larviis  are  six- 
legged.  They  live  on  or  under  the  skin  of  birds  and  mammals,  on  which  they 
produce  the  skin  disease  known  as  scabies,  or  itch. 

[The  Sarcoptid(E  attack  the  hairs,  feathers  or  epidermis  of  birds,  animals  and 
man,  living  as  permanent  parasites.  The  punctures  they  produce  are  followed  by 
the  formation  of  more  or  less  thick  crusts  or  scabs,  beneath  which  the  mites  live  and 
breed  (so  called  scab,  mange  and  itch).  Most  are  oviparous,  some  ovoviviparous. 
The  eggs  are  minute,  ovoid,  with  a  thin  semi-transparent  shell.  They  incubate  in 
a  few  days,  varying  from    two   to   ten  or   eleven,   as  a  rule.     Generally  sarcoptic 


Second       ,, 

„     30 

Third 

»     45 

Fourth       „ 

,,     60 

Fifth 

M     75 

Sixth 

M           90 

SARCOPTID^  517 

diseases  lie  dormant  in  winter  and  revive  in  spring  and  summer  in  man  ;  but  in 
animals  with  long  wool,  such  as  sheep,  they  are  most  active  during  winter,  although 
revival  of  active  reproduction  takes  place  in  spring. 

[Speakmg  generally,  for  the  Sarcoptidce  there  are  three  distinct  stages  in  the 
development  of  the  male,  four  in  the  female,  as  follows : — 

[(i)  The  larva.     In  this  stage  only  three  pairs  of  legs  occur. 

[(2)  The  nymph,  in  which  a  fourth  pair  of  legs  appear,  and  which  thus  approaches 
the  adult ;  but  so  far  no  sexual  organs  occur.  Nymphs  are  of  two  sizes — the 
smaller  being  future  miles,  the  larger  females. 

[(3)  The  next  stage  in  the  female  is  the  age  of  puberty^  the  female  now  being 
provided  with  a  vulvo-anal  slit ;  this  so-cdW^di  pubescent  female  is  fertilized  by  the  male. 
The  male  then  dies.    But  the  female  again  casts  her  skin  and  enters  another  stage — 

[(4)  The  ovigerous  female — the  egg-laying  female— which  has  differently  modified 
legs. 

[The  rate  at  which  these  Acari  breed  is  very  great.  Gerlach  has  found  that 
roughly,  in  each  Sarcopt  gallery,  a  female  produces  fifteen  individuals — ten  femailes 
and  five  males — and  that  the  progeny  reproduce  again  in  fifteen  days.  The  table 
given  below  thus  shows  that  one  pair  may  produce  the  enormous  number  of 
1,500,000  descendants  in  three  months  : — 

First  generation  after  15  days      ...                  10  females  ...  5  males 

100        ,,  ...  50  ,, 

1,000        ,,  ...  500  ,, 

10,000        ,,  ...  5,000  ,, 

100,000        ,,  ...  50,000  ,, 

...     1,000,000        ,,  ...  500,000  ,, 
:=  ly^oOyOOO  individuals. 

[These  Acaririce  are  divided  into  three  distinct  sub-families,  namely  the  Cyto- 
lichince^  Sarcoptince,  Caiiestriniino'. 

[The  Sarcoptince  2\o\\^  interest  us  here,  and  of  the  nine  genera  the  three  following 
are  the  most  important  : — 

[(i)  Sarcoptes,  Latreille  ;  Eusarcoptes. 

[(2)  Psoroptes,  Gerv.  ;  Dermatodectes,  Gerlach  ;  Dermatocoptes,  Fiirstenberg. 

[(3)  Chorioptes,  Gerv.  ;  Symbiotes,  Gerlach  ;  Dermatophagus,  Fiirst.  ;  Sarco- 
dermatocedes,  Del. 

[The  following  are  the  main  characters  of  these  three  genera:  — 

\Sarcoptes — round  or  slightly  oval;  the  two  posterior  pairs  of  legs  being  nearly  or 
quite  concealed  beneath  the  body;  the  tarsi  end  in  simple  long  pedicles,  with 
ambulatory  suckers. 

{Psoroptes — oval;  the  legs  are  all  visible  outside  the  niargin  of  the  body;  the 
ambulatory  suckers  are  carried  on  long  triangulated  stalks ;  the  male  has  copulatory 
suckers  and  abdominal  prolongations. 

[Chorioptes — oval ;  legs  long,  thick,  all  visible  ;  ambulatory  suckers  very  wide, 
carried  at  the  end  of  simple,  short  pedicles. 

[Sarcoptes  make  channels  or  furrows  beneath  the  epidermis,  and  in  these  the 
female  lays  her  eggs.  This  form  of  acariasis  is  thus  difficult  to  cure.  It  is  the  cause 
of  human  itch  {vide  Sarcoptes  scabiei). 

[Psoroptes  do  not  make  sub-epidermic  galleries  ;  they  live  and  breed  in  colonies 
beneath  crusts  or  scabs  formed  by  the  changes  they  produce  in  their  host's  skin. 
Sheep  scab  is  a  common  type  of  disease  produced  by  Psoroptes.  This  genus  is  of 
little  importance  as  a  parasite  to  man. 

[Chorioptes  live  as  Psoroptes  ;  they  also  do  not  affect  man.  Otodectes,  Can., 
affecting  cats  and  dogs,  and  others  occur,  but  do  not  affect  man  as  far  as  we  know  at 


5i8 


THE   ANIMAL   PARASITES   OF   MAN 


present    ("Demodicidae    und    Sarcoptidae,"    von    Professor    G. 
P.  Kramer,  Das  Tierreich,  1899).— F.  V.  T.] 


Canestrini    und' 


Sub-family.     Sarcoptinae. 

Genus.     Sarcoptes,  Latreille. 

Sarcoptes  scabiei,  de  Geer,  1778. 

Syn. :  Acarus  scabiei^  de  Geer,  1778  ;  A.psoricus^  Pallas,  1760  ;  A.  siro^  L.,  1778  ;; 
Sarcoptes  exulcerans,  ?  Linn,,  1758,  Nitsch,  1818  ;  S.  hommis.  Raspail,  1834,  and 
Hering,  1838  ;  S.galei,  Owen,  1853  ;  5.  communis^  Delaf.  et  Bourg.,  1862;  S.  scabiei 
\2iX.  hominis^  Megnin,  1880. 

The  body  is  oval  or  nearly  circular  and  whitish  in  colour,  with  trans- 
verse rovN^s  of  striae  partly  interrupted  on  the  back.  There  are  trans- 
verse rows  of  small  bristles 
on  the  dorsal  surface,  and 
groups  of  trichomae  on 
the  front,  sides  and  back. 
There  are  chitinous  hairs  at 
the  base  of  the  legs  ;  the  two 
first  pairs  are  provided  with 
pedunculated  ambulacra  in 
both  sexes,  the  two  posterior 
pairs  terminate  each  with  a 
long  bristle  in  the  female  ; 
in  the  male  the  third  pair 
of  legs  terminate  in  a  bristle, 
the  fourth  pair  with  a  pedun- 
culated ambulacrum.  The 
anus  is  situated  at  the  pos- 
terior border  of  the  dorsal 
surface. 

At  one  time  numerous  species 
were  dififerentiated,  according  ta 
the  form  of  the  Acarus,  tlie  num- 
ber, position  and  size  of  the  hairs 
and  spines,  even  according  to  the 
hosts,  etc.  All  these  characteris- 
tics, however,  fluctuate  so  con- 
siderably that  absolute  differentiation  is  impossible;  the  supposed  species  may  be 
regarded  in  the  same  light  as  Megnin  did,  as  varieties.  It  is  also  hardly  possible 
to  distinguish  the  mite  of  human  scabies  {S.  hominis)  from  that  of  a  number  of 
domestic  animals  {S.  squamiferus).  It  is  best,  therefore,  to  accept  one  single  species 
{S.  scabiei)^  which  may  give  rise  to  different  races  or  castes  by  living  in  the  skin 
of  man  and  mammals,  but  can  pass  from  one  host  to  the  other. 

[Canestrini  and  Kramer,  in  their  monograph  of  the  SarcoptidcCy 
enumerate  eighteen  distinct  species  of  this  genus,  from  the  dog,  goat. 


Fig.  367. — Sarcoptes  scabiei :  female,  dorsal  aspect. 
200/1.     (After  Furstenberg.) 


SARCOPTES   SCABIEI 


519 


•camel,  horse,  ferret,  lion,  wolf,  sheep,  pig,  etc.,  and  two  species 
parasites  of  man  (scabiel  and  scabiei-criistoscv).  There  is  no  doubt 
that  they  are  distinct  species. — F.  V.  T.] 

The  S.  scabiel  of  man  (S.  scabiei  var.  Iwminis)  (length  of  male 
0-2  to  o'3  mm.,  and  breadth  0*145  ^^  0*190  mm.  ;  length  of  female 
c*33  to  o'45  mm.,  and  breadth  0*25  to  0*35  mm.)  lives  in  the  tunnels 
that  it  excavates  in  the  epidermis,  and  attacks  by  preference  places 
with  thin  skin,  such  as  between  the  fingers,  in  the  bend  of  the 
elbows  and  knees,  in  the  inguinal  region,  on  the  penis,  on  the 
mammae,  but  may  also  affect  other  parts.  The  tunnels,  which 
vary  from  a  few  millimetres  to  a  centimetre  and  more  long,  do  not 
run  straight,  but  are  somewhat 
tortuous  ;  the  female  is  found 
at  the  terminal  end.  The  tun- 
nels contain  the  excrement  and 
oval  eggs  (0-14  mm.  in  length) 
of  the  parasite ;  the  males  are 
rarely  met  with,  as  they  die  off 
after  copulation ;  the  females 
die  after  depositing  their  eggs. 
The  six-legged  larvae  hatch 
out  after  four  to  eight  days, 
and  after  about  a  fortnight, 
duringwhich  time  they  change 
their  skins  three  times  and 
undergo  metamorphosis,  they 
begin  themselves  to  burrow. 
Transmission  from  person  to 
person  rarely  is  effected 
through  linen,  but  by  direct 
contact  (as  in  coitus)  ;  trans- 
mission can  be  artificially 
effected  on  horses,  dogs  and 
monkeys,  but  not  on  cats. 

The  smaller  S.  scablei-cnis- 
toscv,  Fiirstenberg,   is   the    cause    of  the  itch    that  occurs   chiefly  in 
Norway ;    it    is    not    certain    whether   this    is  a  distinct   species   of 
itch  mite. 

[This  is  quite  a  distinct  species,  which  is  recorded  from  Germany  and 
France.  Megnin  {Parasitology ^  1880,  p.  165)  described  this  as  S.  scablel 
var.  lupl.  The  female  is  140  fx  long,  340  ^i  broad  ;  the  male  is  170  yit  long 
by  150  fji  broad.  In  Science  (March  3,  1893,  p.  125)  is  recorded  that 
at  the  Indiana  Academv  of  Science  Dr.  Robert  Hessler  referred  to 
**  a   case    of   that   extremely    rare  and    almost    extinct   form   of   itch 


Fig 


-Sarcoples  scabiei  :  male,  ventral  aspect. 
203/I.     (After  Fiirstenberg.) 


520 


THE   ANIMAL   PARASITES   OF   MAN 


known  as  'Norway  itch,'  the  scabies  norvcgica  of  Hebra,  1852. 
The  afflicted  man  was  covered  with  thick,  creamy  white,  leathery 
scales ;  some  of  these  scales  measured  over  an  inch  in  diameter  and 
JL  in.  thick.  A  constant  shedding  of  scales  went  on,  a  handful 
being  gathered  daily.  They  were  found  full  of  mites  and  eggs  and 
riddled  with  passages.  Under  treatment  the  mites  were  killed  and 
the  skin  became  normal.  Dr.  Hessler  made  a  calculation  of  the 
number  of  eggs  and  mites,  amounting  to  ova  and  shells  7,004,000, 
mites  in  all  stages  2,009,000. — F.  V.  T.] 

The  following  forms  may  be  transmitted  from  domestic  animals 
to  man  : — 

(i)  S,  scabiei  var.  eqiii.  Male,  o'2  to  0-23  mm.  long,  0*16  to  0*17  mm.  broad. 
Female,  0*40  to  0*42  mm.  long,  0*28  to  o'32  mm.  broad.     The  horse  is  the  normal  host. 

(2)  S.  scabiei  var.  ovis.  Male,  0*22  mm.  long,  o"i6  mm.  broad.  Females, 
0*32  to  o*44mm.  long,  0*24  to  0*36  mm.  broad.  This  mite  lives  on  sheep,  and  passes 
over  to  goats  and  human  beings;  it  may  also  be  artificially  transferred  to  horses, 
oxen  and  dogs.^ 

(3)  S.  scabiei  var.  caprcc.  Male,  0-24  mm.  long,  o'lSS  mm.  broad.  Female, 
o'345  mm.  long,  0*342  mm.  broad.  On  goats,  passing  from  them  to  horse,  ox,  sheep, 
pig  and  man.  On  the  latter,  in  contradistinction  to  the  varieties  (i)  and  (2),  it  pro- 
duces a  severe  affection. 

(4)  S.  scabiei  var.  caineli.  Frequently  observed  in  man,  chiefly  in  Africa.  A 
few  cases  have  been  observed  in  Europe ;  the  affection  induced  by  it  is  severe. 

(5)  S.  scabiei  vviY.  auchefticr.  Male,  0*245  ^'^^'  ^o^R'  o"i82  mm.  broad.  Female, 
0*34  mm.  long,  0264  mm.  broad.  It  lives  on  the  llama,  and  may  be  transmitted 
to  man. 

(6)  S.  scabiei  var.  suis,  Male,  0*25  to  0*35  mm.  long,  o"i9  to  0*3  mm.  broad. 
Female,  0*4  to  0*5  mm.  long,  0*3  to  0*39  mm.  broad.  In  the  domestic  pig  and  wild 
boar;  occasionally  also  in  man.  The  settlement,  however,  is  usually  of  short 
duration. 

(7)  S.  scabiei  var.  canis.  Male,  0*19  to  0*23  mm.  long,  0*14  to  0*17  mm.  broad. 
Female,  0*29  to  o'38  mm.  long,  0*23  to  0*28  mm.  broad.  In  the  house-dog,  and  also, 
not  unusually,  in  human  beings. 

(8)  and  (9)  .S*.  scabiei  var.  vuipis  and  S.  scabiei  var.  leofiis  of  the  fox  and  lion 
have  likewise  been  observed  on  man. 

These  are  all  distinct  species  and  should  read  as  follows  :  S.  canis,  Gerl. ;  S.  ovis, 
Megn.  ;  S.  egtii,  Gerl. ;  .S".  dromedarii,  Gerv.  {cameli,  Megn.)  ;  S.  auchenice,  Raill.  ; 
S.  suis,  Gerl.  :  S.  vuipis,  Fiirst.  ;  S.  leonis.  Can. 

Sarcoptes  minor,  Furstenberg,  1861. 

Anus  situated  on  the  back,  legs  short,  pedunculated  ambulacra 
broad  ;  living  on  cats  (S.  minor  var.  cati)  and  rabbits  (S.  minor  var. 
cunicnli).  In  cats  this  mite  usually  lives  in  the  cervical  region,  and 
thence  spreads  to  the  ears  and  head  ;  it  usually  causes  the  death  of 
the  infected  animals;  it  is  easily  transferable  from  cat  to  cat,  is 
difficult  to  transmit  to  rabbits,  but  once  settled  on  them  can  easily 

'  [This  mite  produces  the  so-called  '*  black  muzzle  "  of  sheep.— F.  V.  T.] 


SARCOPTES   MINOR 


521 


infect  other  rabbits.  On  the  other  hand,  the  transmission  of  the 
itch  mite  of  the  rabbit  to  the  cat  does  not  succeed.  In  man  S.  minor 
induces  an  eruption  that  disappears  after  about  a  fortnight. 

[S.  minor,  Fiirstenberg,  i86t  ('' Kratzm.,"  viii,  p.  218),  comes  in 
Railhet's  sub-genus  Notoedres,  1893  {''  ZooL,"  ed.  2,  p.  660).  Canes- 
trini  raised  this  to  generic  rank  in  1894  (Pros/).  Acarof.,\\,  p.  724). 

[There  are  three  species  :  (i)  iV.  notoedres,  Mcgnin  =  Sarcoptes 
alepis,  RaiUiet  and  Lucet  {Compt.  rend.  Soc.  de  BioL,  1893,  xlv,  p.  404), 
and  Sarcoptes  notoedres  var.  mnris,  Megnin  {Parasitology,  1880, 
pp.  172-174).  This  occurs  on  the  black  and  brown  rats  and  the 
water-vole. 

[(2)  N.  cati,  Hering,  1838  {N.  acta.  ac.  Leop.,  ii,  18,  xliv,  p.  605, 
figs.  9  and  10),  =  Sarcoptes  minor,  Fiirstenberg  ("  Kratzm.,"  1861,  viii, 
p.  215).     Found  on  the  cat  in  Germany,  France,  Italy,  and  Britain. 

[(3)  iV.  cnniculi,  Gerlach,  1857,  ^'Kratzm.,"  iii,  figs.  20,  21.  It 
lives  on  the  rabbit  and  is  found  in  Germany  and  France. — F.  V.  T.] 


Fig.  369. — Sarcoptes  minor  var.  call :  on  the  left,  female  (lying  on  its 
abdomen) ;   on  the  right,  male  (lying  on  its  back).     (After  RaiUiet.) 

The  itch  mites  of  domestic  animals,  which  belong  to  the  genera  Psoroptes 
(=  Dermatodectes  =  Dermatocoptes)  and  Chorioptes  (Symbiotes  =  Dermatophagus), 
as  a  rule  do  not  infest  and  live  on  man,  even  when  artificially  transmitted.  It  is, 
however,  possible  for  this  to  occur.  Moniez  ("Traite  de  par.,"  1896,  p.  559) 
mentions  that  a  species  of  Chorioptes — probably  C^.  bovis — had  been  found  on  man, 
as  had  also  Demodex  folliculorum.  This  author  also  includes  Derniatophagoides 
scherevieteiuskyi,  Bogdanofif  {Bull.  soc.  imp.  d.  natural.,  Moscou,  1864,  xxxvii, 
p.  341),  which  has  repeatedly  been  found  on  man  in  Moscow  and  Leipzig  (Zurn, 
Ber.  d.  ined.  Ges.,  Leipzig,  1S77,  p.  38),  as  Chorioptes  bovis. 

Other  References  to  Scabies  crustoscz  and  norvegica,  etc. 

(1)  "Ein  P'all  von  Scabies  crustosa  norvegica,'"  Wiirzb.  med.  Zeitschr.,  1,  pp.  134-139, 
pi.  3,  H.  Bamberger. 

(2)  "  Ueber  die  Kratzmilbe  {Acarus  scabiei),'"  Notiz.  a.  d.  Geb.  d.  Nat.  u.  Heilk.,  Weimar 
(1913),  xlii  (11),  Oct.,  pp.  161-166  (1834),  de  Blainville. 

(3)  "  Rapport  sur  le  ciron  de  la  gale  {Acariis  scabiei),''^  Ann.  de  Mus.  d' Hist,  nat.y 
1831  ;  Parasitology,  iv,  pp.  213-232,  dc  Blainville. 


522 


THE   ANIMAL   PARASITES   OF   MAN 


Family.     Demodlcidae  (Mites  of  the  Hair-follicles). 

Small  Acarina,  elongated  in  worm-like  fashion,  with  annulated  abdomen,  and 
without  eyes  or  tracheae.  The  mouth  parts  consist  of  a  suctorial  proboscis  and 
three-jointed  palpi  ;  the  legs  are  short,  and  have  three  segments  with  small  terminal 
ungues.  The  anus  is  situated  on  the  anterior  border  of  the  abdomen  ;  oviparous  ; 
the  larvae  have  six  stumpy  legs.     These  mites  live  in  the  hair-follicles  of  mammals. 

Genus.     Demodex,  Owen. 
Demodex  folliculorum,  Simon,  1842. 


Syn. :  Acarus  folliculoriini^  Sim.,  ] 
Macrogaster  platypus^  Miescher,  1843; 
Steatozoon  folliculorum^  Wilson,   1847. 


843;  Demodex  folliculorum^  Owen,   1843; 
Simonea  folliculoi^um^   P.  Gervais,   1844  ; 


As  in  Sarcoptes  sccibiei,  numerous  varieties  of  this  species  are 
known  ;  the  form  parasitic  on  man  lives  in  the  hair-follicles,  the 
meibomian  and  sebaceous  glands,  and  hardly  ever  causes  incon- 
venience; the  male  measures  o'3  mm.  in  length  and 
the  female  about  0*4  mm.  in  length.  The  eggs 
o'o6  to  o'o8  mm.  in  length,  0*04  to  0*05  mm.  in 
breadth,  and  are  thin-shelled.  The  creatures  are 
always  attached  with  the  head  end  downwards  in 
the  parts  mentioned  ;  they  are  most  frequent  in 
the  sebaceous  glands  of  the  face,  by  the  nose,  lips 
and  forehead,  but  they  may  be  present  on  the 
abdomen  and  on  other  parts  of  the  body.  They 
may  occasionally  obstruct  the  excretory  gland  ducts, 
thus  causing  infiammation  of  the  gland  (comedones) ; 
their  agglomeration  in  the  meibomian  glands  sets 
up  inflammation  of  the  margins  of  the  eyelids. 
There  are  generally  only  a  few  specimens  in  a  gland. 
According  to  some  statements  Demodex  occurs  in 
50  per  cent,  of  mankind  and  even  in  children  ;  they 
survive  the  death  of  their  hosts  by  several  days. 


Fig.  2>1o.  — Demo- 
dex follictilorum  of 
the  dog.  (After 
Megnin.) 


The  variety  living  in  the  dog  {D.  folliculorum  var.  canis) 

is  smaller  than  the  variety  living  in  man,  and  produces  a  skin 

disease  resembling  scabies   in  these  animals.     According  to 

Ziirn  they  may  also  live  on  man  ;   nevertheless,  no  other  investigator  has  recorded 

a  similar  observation,  and  attempts  at  artificial  infection  have  proved  negative.^ 

[Ten  distinct  species  of  Demodex  are  given  by  Canestrini  and 
Kramer  ("  Demodicidae  und  Sarcoptidae,"  Das  Ticrreich,  1899,  vii). 
The  species  are  certainly  distinct. 

[The  species  living  on  the  dog  (D.  canis,  Leydig,  1844)  is  cosmo- 
politan.   According  to  the  British  Medical  Journal  (February  22,  1913, 


'  [This  mite  causes  what  we  know  in  England  as  red  mange  in  dogs.  — F.  V.  T.] 


PENTASTOMIDA  523 

p.  407),  dog  mange  may  be  caught  by  humans.  Whittield  and 
Hobday  describe  in  the  Veterinary  Journal  seventeen  cases  which 
have  come  under  their  observation. — F.  V.  T.] 

Order.     Pentastomida. 
Family.      Linguatulidae. 

Aradmida  grfeatly  altered  in  consequence  of  their  parasitic  manner  of  life  ; 
for  a  long  time  they  were  regarded  as  helminthes.  The  body  is  elongated, 
vermiform,  flattened  or  cylindrical,  and  more  or  less  distinctly  annulated.  The 
head,  thorax,  and  abdomen  are  not  defined  from  each  other  (fig.  371).  The  elliptical 
mouth,  surrounded  by  a  chitinous  ring,  is  situated  at  the  anterior  end,  on  the 
ventral  surface,  and  the  intestine  leading  straight  through  the  body  opens  at  the 
posterior  end.  Two  retractile  hooks  are  at  the  sides  of  the  mouth  (fig.  372)  ;  these 
are  usually  considered  to  be  the  terminal  joints  of  two  pairs  of  legs,  but  it  appears 
to  be  more  correct  to  regard  them  as  the  remains  of  the  antenna?  and  palpi  (Stiles). 
According  to  this  opinion,  the  legs  in  the  adult  state  are  completely  degenerated. 

The  nervous  system  is  reduced  to  an  oesophageal  ring.  No  organs  of  sense  are 
recognizable  except  the  papillae  at  the  anterior  end.  There  are  neither  organs  of 
circulation  nor  of  respiration.^ 

The  sexes  are  distinct.  In  the  small  male  the  sexual  orifice  is  situated  ventrally 
in  the  anterior  part  of  the  body  ;  in  the  female  it  is  placed  near  the  anus.  The 
Linguatulidcc  lay  eggs,  and  from  each  ^%%^  after  being  conveyed  into  an  inter- 
mediate host,  a  four-legged  larva,  with  rudimentary  mouth  parts,  hatches  out.  It 
goes  through  a  series  of  metamorphoses,  and  passes  through  a  second  larval 
condition,  which,  however,  possesses  the  essential  characteristics  of  the  fully 
developed  form.  Sooner  or  later  it  migrates  during  this  stage,  and  reaches  its  final 
host,  mammal  or  reptile,  in  the  nostrils  or  lungs  of  which  the  adult  Linguatulidce 
live.  I 

[As  adults  they  live  as  internal  blood  feeders  in  various  birds, 
reptiles  and  mammals,  especially  in  the  nasal  and  respiratory- 
passages.  The  larval  stage  occurs  in  another  host  in  an  encysted 
condition  ;  this  host  is  usually  an  animal  preyed  upon  by  the  species 
in  which  the  sexual  forms  are  found.  The  larvae  bore  through  the 
walls  of  the  host's  stomach  and  enter  liver  and  spleen  or  brain,  where 
they  encyst ;  here  they  grow  until  they  assume  almost  the  appearance 
of  the  adult.  These  encysted  larvae  on  being  eaten  later  make  their 
way  into  the  nasal  passages  and  lungs,  where  they  mature.  Both 
adults  and  larvae  occur  in  man,  as  mentioned  later. 

[Three  genera  are  recognized  in  this  family  : — 

[(i)  Lingiiafula. — Body  flat,  annulated.     Adults  live  in  the  nasal  sinus. 
[(2)  Porocephalus. — Body  cylindrical,  elongate,  with  often  deeply  cut  rings.    Adult 
in  respiratory  organs  of  snakes,  larvce  in  animals  and  man. 

[(3)  Reighardia.—Q,y\\ndx\Q.d\^  but  not  ringed.     Not  found  in  humans. — F.  V.  T.] 

'  What  are  designated  as  stigmata  in  the  Linguatulides  are  the  orifices  of  sebaceous 
glands. 

33 


524 


THE   ANIMAL   PARASITES   OF   MAN 


Genus.     Linguatula,  Frohlich. 
Linguatula    rhinaria,    Pilger,    1802. 

Syn.  :   Tcenia  rhinaria^  Pilger,  1802  ;  Polysto7na  tcrmoides^  Rud.,  18 10; 
Linguatula  tcrnioides,  Lam.,  1816  ;  Pentastoma  ia-nioides^  Rud.,  18 19. 

The  male  is  white  in  colour,  18  to  20  mm.  in  length,  anterior 
portion  3  to  4  mm.  in  breadth,  posterior  part  0-5  mm.  in  breadth. 
The  female  is  of  a  yellowish  colour,  8,  10,  or  13  cm.  long, 
anterior  part  8  to  10  mm.  and  posterior  part  2  mm.  wide.  The 
brownish  eggs  can  be  seen  in  the  median  line.  The  body  is 
elongated,  rather  flat,  and  exhibits  about  ninety  rings 
or  segments  with  crenellated  borders.  The  hooks 
round  the  mouth  are  strongly  curved  and  are  articu- 
lated to  a  basilar  support.  Eggs  oval,  0*09 /a  in  length, 
0*07  yLt  in  breadth. 

L.  rhinariay  in  the  adult  con- 
dition, lives  in  the  nasal  cavity 
and  frontal  sinus  of  the  dog, 
wolf,  fox,  horse,  goat,  and 
occasionally  of  man  ;  it  causes 
severe  catarrh,  epistaxis  and 
suppuration. 

Development.  —  The  ova, 
which  are  found  in  masses  in 
the  nasal  mucus,  already  possess 
an  embryo  ;  they  are  expelled 
with  the  nasal  secretion,  and 
are  swallowed  by  herbivorous 
mammals  with  their  food, 
mostly  by  hares  and  rabbits, 
but  also  by  sheep,  goats,  oxen, 
horses,  antelopes,  fallow^  deer, 
pigs,  cats,  and  occasionally  also 
by  human  beings.  The  young 
larvae  hatch  out  in  the  stomach; 
they  possess  a  thickened  an- 
terior body  with  rudimentary 
mouth  parts  and  two  pairs  of 
limbs  ;  the  body  gradually  tapers  to  a  short  tail. 

The  larv^  of  the  Linguatulidcv  bore  through  the  intestinal  wall 
and  reach  the  liver,  more  rarely  the  mesenteric  glands,  etc. ;  they 
Here  become  encysted  and  enter  a  sort  of  pupal  stage  in  which  they 
lose  their  limbs  ;  after  several  moultings  and  gradual  growth  the 
second  larval  stage,  having  the  appearance  of  the  adult  Linguatula, 


Fig.  ^71.— Lingua- 
tula rhinaria :  fe- 
male.   Natural  size. 


Fig.  372.— Larva  of 
Linguatula  rhinaria 
{Pentastoma  denticula- 
tum).  Enlarged.  (After 
Leuckart.) 


LINGUATULA   RHINARIA 


525 


sets  in.  About  five  to  six  months  after  infection  the  creatures  have 
become  4  to  6  mm.  long,  possess  eighty  to  ninety  rings,  which  have 
a  series  of  fine  points  on  their  posterior  border;  the  mouth  and 
intestine  are  formed,  the  sexual  organs  mature  and  the  two  pairs  of 
hooks  are  near  the  mouth.  This  larval  stage  (fig.  372)  has  been 
known  for  a  long  time,  but  it  was  regarded  as  an  independent  species 
of  animal,  and  therefore  had  a  separate  name  {Liiigitafiila  scrrata, 
Fr.  ;  Pentastonia  denticnlatiun,  Rud.,  etc.). 

Later  these  Linguatula  larvae  make  an  attempt  to  escape  from  their 
hosts,  and  this,  of  course,  can  only  be  effected  by  means  of  an  active 
migration  ;  they  leave  the  cysts,  and  according  to  their  respective 
positions  in  the  abdominal  or  pleural  cavities  they  reach  the  bronchi 


Fig.  373. — Linguatula  rhinaria:  on  left,  eggs  in  gelatinous  covering,     iio/i. 
On  right,  first  larval  stage.     300/1.     (After  M.  Koch.) 


or  the  intestine,  and  finally  pass  out ;  they  may  be  again  sniffed  up 
by  dogs  and  settle  in  their  nasal  cavities.  Still  this  outward  migra- 
tion does  not  appear  to  be  necessary  for  further  development.  A 
portion  of  the  larvae  gain  access  to  the  nasal  cavities  directly  through 
the  trachea,  and  thus  herbivorous  mammals  certainly  become  directly 
infected.  In  most  cases  the  infection  of  dogs,  wolves  and  foxes,  that 
is,  of  carnivorous  mammals,  takes  place  through  consuming  the 
bodies  of  mammals,  or  parts  of  them,  such  as  the  liver  and 
lungs,  which  are  alTected  with  the  second  larval  form  ;  in  any  case 
most  larvae  obtain  access  first  to  the  stomach  of  their  host,  from 
here  they  make  an.  active  migration  through  the  oesophagus  to  the 
oral  and  nasal  cavities,  in  which  they  settle.     It  is  possible  also  that 


^26  THE   ANIMAL   PARASITKS   OF   MAN 

the  same  larvae  which  are  free  in  the  oral  cavity  when  the  food  is 
being  eaten  migrate  into  the  nasal  cavities.  After  being  stationary 
a  fresh  skin  is  formed  and  the  spine-bearing  cuticula  are  thrown  off. 
The  male  attains  its  full  size  in  the  fourth,  and  the  female  in  the  sixth 
month.  The  duration  of  life  is  stated  to  be  from  fifteen  months  to 
several  years. 

L.  rhinaria  has  been  observed  in  man  in  the  adult  as  well  as  in  the  larval 
condition  {Pentastoma  denticulatum).  Zenker  first  called  attention  to  the  occurrence 
of  the  larva  in  man,  having  found  it  nine  times  in  the  liver  in  i68  autopsies.  Heschl 
found  it  twice  in  Vienna  in  twenty  autopsies,  Virchow  found  it  in  Wiirzburg  and  Berlin, 
Wagner  in  Leipzig  (lo  per  cent.),  and  Frerichs  in  Breslau  five  times  in  forty-seven 
autopsies.  The  parasite  is  nmch  less  frequent  in  Switzerland.  According  to  Klebs, 
one  case  occurs  in  900  autopsies,  and  according  to  Zaeslin  two  cases  occurred  in  Basle 
to  1,914  autopsies.  In  the  Seamen's  Hospital  in  Kronstsidt  P.  den^i'cu/a/i^m  has  been 
found  six  times  in  659  autopsies.  It  was  almost  always  the  liver  that  contained  one 
or  a  few  specimens.  The  parasite  was  very  rarely  found  in  the  kidney  or  spleen,  or 
encysted  in  the  intestinal  wall.  The  adult  L.  rhinaria  is  far  more  rarely  observed 
in  man. 

A  case  reported  by  Landon  that  related  to  a  blacksmith  of  Elbing  is  particularly 
interesting.  This  man  accompanied  the  campaign  of  1870;  he  soon,  however,  fell 
ill  with  pains  in  the  liver,  accompanied  by  icterus  and  intestinal  disorders.  Soon 
after  the  war,  and  after  the  symptoms  were  reduced  to  icterus  and  weakness,  bleed- 
ing of  the  nose  set  in  and  continued  with  slight  intermissions  for  seven  years  ;  an 
unpleasant  sensation  of  pressure  in  the  left  nasal  cavity  set  in,  with  inflammatory 
swelling  of  the  mucous  membrane.  At  last,  in  the  summer  of  1878,  when  the 
pressure  in  the  nose  had  considerably  increased,  a  Linguatula  was  expelled  from  the 
nose  with  a  violent  attack  of  sneezing,  and  lived  for  three  days  longer  in  water.  The 
bleeding  of  the  nose  then  ceased  and  the  patient  soon  recovered.  There  can  be  no 
doubt  that  the  first  illness  was  connected  with  the  invasion  in  the  liver  of  numerous 
larvae  of  Pentastoma,  and  disappeared  after  their  encystment  ;  one  or  a  few  of  these 
must  subsequently  have  found  its  way  to  the  nose  and  settled  there. 

Genus.     Porocephalus. 
Porocephalus  constrictus,  v.  Siebold,  1852. 

•    Syn.  :  Nematoideutn  honmiis^  Diesing,  185 1  ;  Pentastomuni  constrictuin^ 
V.  Sieb.,  1852  ;  Porocephalus  constrictus,  Stiles,  1893. 

Porocephalus  is  distinguished  from  t  inguatula  by  its  cylindrical 
body  and  by  certain  internal  structures.  Porocephalus  constrictus 
is  at  present  only  known  in  its  larval  stage.  It  is  milk  white  in 
colour  with  golden-yellow  booklets.  Number  of  rings,  twenty-three. 
Length  13  mm.,  breadth  2*2  mm.  There  are  no  prickles  on  the 
posterior  border  of  the  annulations  of  the  body. 

This  species  was  first  discovered  by  Pruner  encysted  in  the  livers  of  two 
negroes  in  Cairo.  Bilharz  reported  two  further  cases  in  which  the  parasites 
were  encysted  in  the  liver  and  in  the  mucosa  of  the  intestine  ;  a  few  other  observa- 
tions have  been  made  by  Fenger,  Aitken,  Giard  and  Chalmas.  Aitken's  report  deals 
with  soldiers  of  the  British  Colonies  in  Africa.     The  parasites  were  discovered  in 


POROCEPHALUS   CONSTRICTUS  527 

the  liver  as  well  as  in  the  lung,  and  appear  to  have  been  the  cause  of  death  in  one 
case  (pneumonia,  peritonitis), 

Pruner  has  found  the  same  parasite  also  in  the  Hver  of  the  giraffe. 

It  has  recently  been  assumed  that  Porocephaliis  constrictus  is  the  larva  of 
Pentasto7na  fnoniliforme,  Diesing,  1835,  that  attains  a  length  of  70  mm.  and  lives 
in  the  lungs  of  African  Pythonides.  The  larva  is  known  to  have  been  ejected 
from  monkeys  {Cercopithecus  albo ocularis ^  Cynocephaliis  maifnon)^  from  the  giraffe 
{Cajnelopardalis  ^iraffd)^  from  a  species  of  hyaena  {Proteles  cristatus)^  and  should 
be  expected  to  occur  frequently  in  smaller  mammals  which  have  been  swallowed 
by  African  serpents  of  enormous  size. 

[The  three  species  of  Pcntastoniidce,  or  tongue  worms,  found  in 
man  are  LinguaUtla  serrata,  FroHch  ;  Porocephaliis  aruiillatuSy  Wyman  ; 
and  Pentastoma  iiionilifon/iis,  Diesing. 

[(i)  Lingiiatiila  serrata  has  been  referred  to  under  a  great  number 
of  names. ^  It  is  a  frequent  parasite  in  dogs,  oxen  and  sheep ;  as  an 
aduh  in  the  dog  and  also  in  the  fox  and  wolf.  The  nymphal  stage  is 
found  in  rats,  hares,  rabbits,  the  horse,  oxen,  sheep,  goats,  pigs,  camels, 
deer,  the  African  and  long-eared  hedgehogs,  porcupine,  guinea-pig 
and  peccary.  In  man  it  is  found  in  both  adult  and  nymphal  stages. 
Sambon  says  the  nymphal  stage  is  of  frequent  occurrence,  but  is 
usually  overlooked.  Zenker,  who  first  found  it  in  man,  obtained  it  in 
nine  out  of  160  post-morteuis,  usually  encysted  in  the  liver.  It  is  then 
said  to  be  harmless.  Landon,  in  1878,  found  the  adult  in  man,  the 
patient  suffering  from  epistaxis  for  about  seven  years ;  in  the  end 
during  a  fit  of  sneezing  the  living  parasite  was  ejected  through  the 
nostril.  This  case  is  of  particular  interest  as  it  appears  to  suggest 
that  this  Acarid  may  now  and  then  pass  its  entire  development  in  the 
same  host,  or  at  any  rate  may  actively  migrate  from  the  liver  to  the 
nasal  cavities  after  a  period  of  encystment  in  the  liver  or  elsewhere, 
which  has  also  been  observed  in  herbivorous  animals  {vide  also  p.  526). 

[It  is  recorded  from  man  in  Central  America  (Darling,  Bull. 
Soc.  Path,  exot.,  191 2,  v,  p.  118;  and  again  Arch.  Int.  Med.,  1912,  v, 
p.  401),  also  from  Rio  de  Janeiro  {Mem.  Inst.  Oswaldo  Cruz,  1913, 
fasc.  ii,  p.  125)  by  Faria  and  Travassos. 

[(2)  Porocephaliis  aruiillatus,  Wyman,  is  also  known  under  a  variety 

'  [Synonymy  given  by  Sambon  :  Adult  form,  Tenia  lanceole,  Chabert,  1787;  Ver  rhinaire, 
Chabert,  1787;  Tcenia  rhinaris,  Pilger,  1805;  Tania  lanceolata,  Rudolphi,  1805;  Cochlus 
r/iinariiis,  Rudolphi,  1805;  Prionoderma  rhinaria^  Rudolphi,  1808;  Polyslovia  t(tnioides, 
Rudolphi,  1809;  Linguatula  tcEuioides,  Lamark,  1816  ;  Prionodervia  lanceolata,  Cuvier, 
1817  ;  Pentastoma  ta;moides,  Rudolphi,  1819;  Linguatnla  lanceolata,  de  Blainville,  1828; 
Lingiiatiila  rhinaris,  Railliet,  1885  ;  Linguatnla  caprina,  R.  Blanchard,  1900.  Nymphal 
form  :  Linguatnla  serrata,  Frolich,  1789;  Tcvnia  capraa,  Abildgaard,  1789;  Tcvtiia  caprina^ 
Gmelin,  i8co  ;  Poly  stoma  serrata,  Gueze,  1800  ;  Halysis  caprina,  Zeder,  1803  ;  Linguatula 
denticulata,  Rudolphi,  1805  ;  Echinorhynchus  caprece,  Braun,  1809;  Tetragulus  caprice,  Bosc, 
1810;  Pentastoma  denticulatum,  Rudolphi,  1819 ;  Pentastoma  e/narginatum,  Rudolphi, 
1819;  Pentastofnafera,  Creplin,  1829;  LAnguatulaferox,  Gros,  1849. 


528  THE   ANIMAL   PARASITES   OF   MAN 

of  names.^  This  species  is  widely  spread  over  tropical  Africa.  The 
adult  sta<:^e  is  found  in  pythons  and  puff-adders,  the  nymphal  in  the 
chimpanzee,  Sykes  monkey,  mandrill  and  other  monkeys,  the  lion, 
leopard,  banded  ichneumon,  Aard  wolf,  dog,  black  rat.  South  African 
reedbuck  and  the  giraffe.  The  adult  has  never  been  found  in  man 
or  any  mammal.  No  fewer  than  sixteen  cases  of  the  nymphal  form,. 
Sambon  tells  us,  have  been  found  in  man,  and  it  is  probably  much 
more  widespread  than  at  present  know^n.  So  far  it  has  only  been 
found  in  the  African  natives.  This  species  has  sixteen  to  seventeen 
body  rings  in  the  male,  eighteen  to  twenty-two  in  the  female,  and  the 
body  does  not  taper  as  much  as  in  the  next  species. 

[(3)  Peiiiasionia  uioiiiliforjiiis,  Diesing,'Ms  an  Oriental  species,  found 
in  India,  Indo-Chma  and  South  China,  and  the  Malay  Archipelago. 
The  adult  occurs  in  both  the  Indian  and  reticulated  pythons. 
The  nymphal  stage  has  been  found  in  monkeys,  the  tiger,  the  civet  and 
the  Indian  otter. 

[The  nymph  has  twice  been  found  in  man  ;  in  one  case  in  the  liver 
of  a  Filipino,  the  other  in  the  serous  coat  of  the  small  intestine  of  a 
native  of  Sumatra. 

[This  species  can  be  told  by  the  female  having  twenty-nine  to 
thirty-three  body  rings,  the  male  twenty-six,  and  the  annulations  are 
more  bead-like  and  less  prominent  than  in  the  African  species. 

[In  addition  to  these  three,  Sambon  thinks  it  probable  that  others 
occur  m  man. — F.  V.  T.] 

Other  References  to  Pent astomi dee. 
(i)  "Die  Wanderung  des  Pentastomum  deniiculahim  beim  Rinde,"  Centralbl.  f.  Bakt . 
u.  Parasitenk.,  Jan.  2,  1889,  v  (i),  pp.  1-5,  V.  Bates. 

(2)  "II  Pentastoma  vicnilifor7ne.  Dies.,  nella  pantera,"  Med. -vet.  Torino,  1877,  4  s., 
vi  (12),  pp.  529-532,  R.  Bassi. 

(3)  "  On  the  Organization  and  Development  of  Linguatula  (Pentastoma),  accompanied 
with  the  description  of  a  new  species  from  the  abdominal  cavity  of  the  mandrill,"  Ann.  and 
Mag.  Nat.  Hist.,  1848,  2  s.  ii  (7),  2,  pp.  69-70,  v.  Beneden. 

(4)  "De  la  Linguatula  ferox  [Pentastoma  denticulatum  aut  serratnm),''  Bnll.  Acad.  roy. 
d.  Sci.  d.  Belg.,  1855,  xxii,  pt.  I  (I),  pp.  4-10,  V.  Beneden. 


*  Adult  form  as  Linguatula  armillata,  Wyman,  1847  ;  Pentastomum  poly zonum ,  ITarley, 
1856;  Pentastoftmmarmillatum,  Leuckart,  i860;  Pentastomum  armillatum,  Diesing,  1864; 
Porocephaliis  armillatus.  Stiles,  1893 ;  Porocephalus  polyzonus,  Stiles,  1893  :  Porocephalus 
moniliformis,  Neumann  (in  part),  1899.  Nymphal  form  :  Linguatula  diesingii,  van  Beneden, 
1849;  Pentastomum  euryzomim,  Diesing,  1850;  Nematoideum  hominis,  Diesing,  185 1  ; 
Pentastomum  constrictum,  von  Siebold,  1852  ;  Linguatula  constricta,  Kuchenmeister,  1855  ; 
Pentastoma  leonis,V^e6\.,  1863;  Pentastoma  fornatum,  CohhoXei,  1879;  Pentastomum  protelis, 
Hoyle,  1883  ;  Porocephalus  constrictus.  Stiles,  1893  5  Linguatula  constrictor,  Galli-Valerio, 
1896;  Pentastomum  diesingii,  Shipley,  1898. 

2  The  synonomy  is  as  follows  :— Adult  form  :  Pentastoma  moniliforme,  Diesing,  1835  ^ 
Linguatule  mofiiliforme,  Megnin,  1880  ;  Porocephalus  moniliformis,  Stiles.  Nymphal  form  : 
Pentastoma  fornatum,  Creplin  (in  part),  1849  ;  Pentastoma  wedlii,  Cobbold,  1866  ;  Penta- 
stoma aonycis,  Macalister,  1874  ;  Porocephalus  armillatus.  Stiles  (in  part),  1908. 


INSECTA 


529 


(5)  "Note  sur  quelques  pentastomes,"  Btill.  Acad.   roy.  d.   Set.  de  Belg.,    1857,   26, 
2  s.,  ii  (5),  pp.  29-30,  V.  Beneden. 

(6)  "  Ueber  das  Petitastoma  in  de  gekrosdrusen  den  Schafe,"  Repert.  d.  Thierh.  Stuttg.^ 
1861,  xxii,  pp.  37-38,  Collin. 

(7)  "  Eine  Linguatula  aus  der  Mesenterialdriise  des  Schafes  und  Dromedars  als  zweites 
ungesche.  Stadium  von  Pent,  taenioides ,''  Notiz.  ti.  Tagsber.  11.  d.  Geb.  d.  Nat,  u.  Heilk. 
Jena,  1862,  v,  pp.  127,  128,  Colin. 

B.     INSECTA  (Hexapoda). 

Three  separate  regions  can  always  be  distinguished  in  the  body  of  insects, 
namely,  the  head,  thorax  and  abdomen.  The  head  is  a  roundish  unsegmented 
capsule  and  possesses  four  pairs  of  appendages.  The  first  pair  are  the  various 
shaped  feelers  (antennae),  which  are  placed  on  the  superior  surface  of  the  head  next 
to  the  eyes  ;  then  more  ventrally  placed  a  pair  of  upper  jaws  (mandibles)  without 
palpi  and  without  articulations  ;  they  are  powerful  masticatory  organs.^  The  first 
pair  of  lower  jaws  (maxilhe)  are  jointed  and  bear  a  palpus  (palpus  maxillaris) ;  the 
second  pair  of  maxillae  are  soldered  together  and  form  the  lower  lip  (labium),  and 
likewise  carry  a  palpus  labialis  on  each  side.  The  upper  lip  (labrum),  as  well  as 
the  other  parts  (which,  however,  are  only  appendages),  belong  to  the  mouth,  which 
IS  really  formed  of  a  number  of  closely  united  pieces.  The  mouth  parts  are  modified 
according  to  the  functions  required  of  them.  Coleoptera,  Neuroptera^  and  Ortho- 
ptera  have  biting  or  masticatory  mouth  parts  which  conform  with  the  scheme 
described  above.  In  the  licking  mouth  parts  of  the  Hymenoptera  the  maxillae  and 
under  lip  are  considerably  elongated,  while  the  mandibles  retain  their  form 
and  are  used  for  triturating  the  food  ;  in  the  Lepidoptera  nearly  all  the  mouth 
parts  are  shortened  except  the  maxillae,  which  form  a  long  and  sometimes  spirally 
rolled  suctorial  proboscis  ;  the  Dipiera  and  Rhynchota  have  piercing  and  sucking 
mouth  parts.  The  mandibles  and  maxillae  are  metamorphosed  into  needle-like 
structures,  while  the  suctorial  apparatus  is  formed  by  the  labrum. 

The  thorax  consists  of  three  segments,  which  are  frequently  united  ;  ventrally  it 
carries  three  pairs  of  legs,  which  consist  of  a  definite  number  of  articulated  pieces 
joined  together.  Their  form  also  changes  according  to  their  function,  so  that  legs 
for  running,  walking,  digging,  swimming,  jumping,  and  preying  are  seen.  A  pair 
of  wings  are  respectively  attached  to  the  last  and  last  but  one  thoracic  rings,  and 
these  may  be  traced  back,  not  to  metamorphosed  appendages,  but  to  tracheal 
branchia.  They  are  composed  of  chitinous  membranes  supported  by  branched 
structures  (veins  or  ribs).  Their  size  and  formation  vary  ;  they  are  seldom  of  equal 
size  and  form  {S europterd) ;  often  the  posterior  wings  are  larger  than  the  anterior 
wings,  the  former  then  only  serving  as  protective  coverings  for  the  latter 
{Coleopterd),  or  the  anterior  wings  are  larger  {Lepidoptera),  or  the  posterior  wings 
are  shortened  or  are  entirely  absent  {Dipiera);  and  finally  there  are  insects  in 
which  both  pairs  of  wings  are  lacking.^ 

The  abdomen  retains  its  segmentation,  but,  with  the  exception  of  a  few  groups 
related  to  the  primitive  forms  of  insects,  has  no  appendages  in  the  imago  condi- 
tion ;  the  abdomen  usually  consists  of  ten  segments,  on  the  last  of  which  the  anus 
is  situated. 

'  [The  mandibles  are  only  powerful  masticatory  organs  in  biting-mouthed  insects 
{Mandibulata) ;  in  the  sucking  or  piercing-mouthed  insects  they  may  be  absent,  or  in  the 
form  of  needle-like  stylets  {Haustellata).—Y .  V.  T.] 

^  [As  in  the  order  Aptera,  which  includes  the  Thysanura  and  Collembola,  and  also 
exceptions  in  other  orders,  as  the  fleas  amongst  Diptera,  the  Mutillus  and  ants  amongst 
Hymenoptera. — F.  V.  T.] 


530  THE   ANIMAL   PARASITES   OF   MAX 

We  need  only  observe  the  following  characters  in  considering  the  anatomy  of 
insects  : — 

The  EPIDERMIS  consists  of  the  chitinous  cuticle,  which  is  separate  from  the 
cellular  layer  beneath  (hypodermis) ;  the  various  appendages  are  supported  by  the 
chitinous  layer. 

The  INTESTINAL  CANAL  usually  consists  of  the  anterior,  median  and  terminal 
intestine,  and  as  a  rule  passes  straight  through  the  body  ;  salivary  glands  discharge 
into  the  anterior  part,  and,  in  some  cases,  yield  a  stiffening  secretion  which  serves 
for  spinning  webs  ;  numerous  or  scanty  hepatic  tubes  are  appended  to  the  median 
intestine,  while  on  the  border  between  the  median  and  terminal  intestine  open  four 
to  six  long  tubes  (vasa  malpighiana),  which  act  as  urinary  organs.  Finally  the  end 
portion  of  the  intestine  carries  various  glands  (anal  and  rectal  glands,  etc.). 

The  CENTRAL  NERVOUS  SYSTEM  agrees  in  structure  with  that  of  the  Annelids, 
but  is  more  highly  developed.  The  pharyngeal  ring  surrounds  the  front  part  of  the 
intestine  ;  the  sensory  nerves  originate  from  its  superior  pharyngeal  ganglia 
and  are  the  seat  of  the  higher  psychical  functions;  the  inferior  pharyngeal 
ganglia  govern  the  mouth  parts,  and  in  addition  appear  to  regulate  the  movements 
(cerebellum)  of  the  vertebrates. 

The  chain  of  GANGLIA  lying  on  the  ventral  side  of  the  abdomen  consists  primi- 
tively of  pairs  of  ganglia  corresponding  with  the  twelve  segments,  which  are  connected 
by  longitudinal  and  transverse  commissures.  But  many  changes  in  the  ganglia  may 
be  seen  in  insects  caused  by  partial  or  entire  amalgamation  of  single  ganglia,  so  that 
in  a  few  cases  only  one  abdominal  ganglion  is  present.  In  conclusion,  a  definite 
INTESTINAL  NERVOUS  SYSTEM  is  always  present. 

Of  the  organs  of  sense  the  FACETTED  EYES,  situated  at  the  sides  of  the  head, 
deserve  special  mention,  as  do  also  the  ORGANS  OF  TOUCH  and  SMELL,  situated  on 
the  antennae,  and  the  organs  of  hearing  and  taste,  or  finer  sensations,  situated  at 
the  mouth  and  in   the  buccal  cavity. 

The  sounds  emitted  by  insects  are,  as  a  rule,  produced  by  the  friction  or  beating 
of  certain  chitinous  parts,  but  sounds  are  also  produced  in  breathing  (flies). 

The  organs  of  respiration,  the  so-called  trachea?,  are  highly  developed  ;  there 
are  openings  (stigmata)  at  the  sides  of  the  body  which  draw  in  air  by  means  of  the 
active  participation  of  the  muscles  of  the  body.  The  number  of  stigmata  varies 
between  two  and  ten  pairs  ;  the  tracheae  themselves  branch  off  from  the  trunks  in 
the  most  varied  manner,  and  carry  air  to  the  internal  organs. 

The  colourless  BLOOD  circulates  between  the  tissues  and  organs,  and  is  kept 
circulating  by  the  contraction  of  a  chambered  dorsal  vessel  provided  with  ostia, 
and  which  terminates  in  a  short  aorta  opening  at  the  anterior  end. 

Insects  are  SEXUALLY  distinct  ;  their  sexual  glands  are  in  pairs  and  have  a 
tubular  structure,  but  the  testicular  tubules  are  united  together  by  a  capsule  into  an 
oval  testicle  ;  exceptionally,  also,  the  excretory  canals  are  double,  as  also  the  sexual 
orifices  ;  usually  the  paired  canals  unite  into  a  single  oviduct  or  spermatic  duct 
which  terminates  at  the  posterior  end  of  the  body  after  receiving  the  products  of 
various  glands. 

As  to  the  history  of  the  development  of  insects,  all  that  is  necessary  to 
mention  here  is  that  the  young  hatched  from  eggs  only  exceptionally  (as  in 
Apterygota)  resemble  the  adult  parent  (insecta  ametabola)  ;  as  a  rule  they  differ 
from  them  not  only  in  the  shape  of  the  body,  but  also  more  or  less  by  their  manner 
of  life,  and  only  attain  the  form  of  the  parent  through  metamorphosis.  This  is  a 
gradual  process  (insecta  hemimetabola)  in  the  Rhy?ichota  and  Orthopiera,  or  a 
sudden  one  with  a  stage  of  inanition  (insecta  metabola)  in  the  other  orders.  This 
stage  of  rest  or  inanition,  the  PUPA,  concludes  the  larval  life  (caterpillar,  maggot, 


INSECTA 


531 


etc.)  ;  during  the  pupal  stage  no  nourishment  at  all  is  taken,  but  the  internal 
organs  undergo  changes  ;  in  some  forms  the  rest  is  not  absolute,  as  voluntary 
local  movements  may  take  place  (pup^  of  gnats). 

The  insects  are  divided  into  numerous  orders  according  to  the  form  of  the 
mouth  parts,  the  structure  of  the  wings,  as  well  as  the  manner  of  the  development ; 
with  the  exception  of  the  lowest  group  {Apterygotd)^  which  is  most  nearly  related  to 
the  ancestors  of  the  insects,  and  which  has  no  wings  and  undergoes  no  metamorphosis, 
all  the  remaining  orders,  which  are  termed  Pterygoia^  have  wings  on  the  thorax, 
though  there  are,  of  course,  a  few  species  and  families  of  this  group  which  have  lost 
their  wings. 

The  Pterygota  include — 

(i)  Orthoptera. — Biting  mouth  parts,  anterior  wings  leathery,  posterior  wings 
thin,  folded  longitudinally  ;  metamorphosis  incomplete  (grasshoppers, 
crickets,  cockroaches). 

(2)  Pseudo7ieuroptera. — Biting  mouth  parts,  wings  of  equal  size,  thin,  not  folded 

up  (dragon-flies,  hair  and  feather  lice,  termites). 

(3)  Rhynchota  or  Hemiptera. —  Mouth  parts  formed  for  puncturing  and  sucking  ; 

wings  alike,  or  the  anterior  wings  may  be  thickened,  parchment-like  at 
their  base  (plant  lice,  cicadae,  bugs  and  true  lice). 

(4)  Neuroptera. — Biting     mouth    parts ;     wings    alike,    thin  ;     metamorphosis 

complete  (ant-lions,  lace-wing  flies,  etc.). 

(5)  Trichoptera. — Licking  mouth  parts  ;  anterior  wings  narrow,  posterior  wings 

longitudinally  folded,  both  ornamented  with  little  hairs  ;  the  larvae  are 
worm-like  in  form,  live  in  water,  and  breathe  through  tracheal  gills 
(may  flies,  etc.). 

(6)  Lepidoptera. — Suctorial  mouth  parts  ;  wings  covered  with  scales  (butterflies). 

(7)  Coleoptera. — Biting  mouth  parts  ;    anterior   wings  thickened    and    differ  in 

colour,  appearance  and  function  from  the  thin,  folded  posterior  wings 
(beetles). 

(8)  Hyinenoptera. — Mouth    parts    for    licking    and    biting  ;    the    wings    alike, 

membranous  (ichneumon  flies,  ants,  wasps,  bees,  humble  bees). 

(9)  Dipiera. — Mouth  parts  formed  for  puncturing,  sucking  or  licking  ;  posterior 

wings  degenerated  (gnats,  flies,  gadflies,  fleas). 

(10)  Strepsiptera. — Anterior  wmgs   shortened  ;    the  female   without  wings    and 

living  parasitically  (fan-wings). 

The  parasites  of  man   occur  amongst   the  Rhynchota^  Coleoptera, 
and  amongst  the  Diptera. 

[The  most  usual  and  recent  classification  of  the  Hexapoda  is  the  following  : — 
(i)    Apte7'a. — Wingless  insects  ;  scarcely  any  metamorphosis. 

(2)  Neuroptera.— Y QMX   membranous    wings,    frequently    with    much    network  ; 

the  front  pair  not  much,  if  at  all,  harder  than  the  under  pair  ;  the  latter 
with  but  little  or  no  fan-like  action  in  closing  ;  mandibulate  ;  meta- 
morphosis variable,  but  rarely  complete. 

(3)  Orthoptera. — Four    wings  ;    front    pair   coriaceous    or    leather-like,    usually 

smaller  than  the  other  pair,  which  are  of  more  delicate  texture  and 
contract  in  repose  like  a  fan  ;    mandibulate  ;    metamorphosis  complete. 

(4)  Thysanoptera. — Four    very    narrow    fringed    wings  ;     mouth     imperfectly 

suctorial  ;  metamorphosis  slight. 

(5)  Hemiptera. — Four  wings  ;  the  front  pair  either  all  transparent  or  with  the 

basal  half  leathery;  mouth  suctorial;  metamorphosis  slight. 


532  THE   ANIMAL   PARASITES   OF   MAN 

(6)  Diptera. — Two  membranous    wings    only  ;    mouth    suctorial,    very   varied  ;; 

metamorphosis  complete. 

(7)  Lepidoptera. —  Four   large    wings    covered    with    scales  ;    mouth    suctoriaU 

metamorphosis  great. 

(8)  Hymenoptera.—Your  membranous  wings  ;  front  pair  larger  than  hind,  which 

do   not    fold   up   in    repose  ;    mandibulate,    sometimes    with    a   tubular 
proboscis ;  metamorphosis  complete. 

(9)  Coleoptera. — Four  wings,  the  front  pair  hard  and  horny  (elytra),  meeting  in 

a  line  over  the  back  and  covering  the  delicate  hind  pair  ;  mandibulate  ; 
metamorphosis  complete. 
[There  are  two  other  well-known  arrangements,  namely,  Packard's  and  Brauer's, 
of  recent  date,  but  the   one   given   here,  which   is  based   on   Linnaeus' 
grouping  by  Dr.  Sharp,  is  by  far  the  simplest. — F.  V.  T.j 


Order.     Rhyncota.^ 

The  lower  lip  forms  a  long  thin  tube  that  can  be  turned  back  (rostrum),  and 
within  which  lie  the  setaceous  mandibles  and  maxillae ;  the  first  thoracic  segment  is 
not  united  with  the  two  posterior  ones  ;  the  anterior  wings  are  usually  leathery  as 
far  as  the  centre. 

{a)    RhYNCOTA   APTEKA    PARASITICA. 

Family.     Pediculidae  (Lice). 

The  lower  lip  is  transformed  into  a  projecting  rostrum  provided  with  barbed 
booklets  in  which  the  hollow  extensile  sucker  (maxillae  and  mandibles)  is  situated  ; 
no  wings ;  no  metamorphosis  ;  only  simple  eyes  ;  the  antennae  are  five-jointed, 
the  feet  possess  hook-like  terminal  structures ;  the  barrel-shaped  eggs  (nits)  are 
deposited  on  the  hair  of  the  host. 

[The  lice  or  PediculidcE  are  also  known  as  Anoplura  and  Siphunculata. 

[They  have  been  split  up  into  a  number  of  families  and  sub-families  and  a 
number  of  genera,  but  as  far  as  this  work  is  concerned  it  is  best  to  retain  the  single 
family  Pediculidcr. 

[Only  the  three  species  mentioned  here  are  common  parasites  of  man,  but  now 
and  then  horse  and  cattle  and  sheep  lice,  Hcematopinus^  may  cause  transitory 
annoyance.— F.  V.  T.] 

Genus.     Pediculus,  Linnaeus. 

Pediculus  capitis,  de  Geer,  1778. 

Male  I  to  1-5  mm.  in  length,  female    1*8  to  2*0  mm.   in  length. 

The  colour  varies  from  light  grey  to  black  according  to  the  colour  of 

the   hair   of   the   human  race   upon  which  they  are  parasitic.     The 

abdomen    has    eight   segments,  of   which   the   six  central   ones  are 

'  [Usually  known  as  Hemiptera,  There  are  two  sub-orders,  Heteroptera  and  Homoptera. 
The  former  have  the  base  of  the  front  wings  coriaceous  ;  the  latter  have  all  four  wings 
membranous.  The  Homoptera  are  Aphides  or  plant  lice  and  scale  insects  {^Coccida),  none 
of  which  attack  man.  Recently  an  interesdng  case  has  been  reported  to  me  where  certain 
Aphides  had  been  passed  in  human  urine.  One  species  was  Rhopalosiphiim  dianihi,  the 
other  found  in  the  urine  was  the  hop  aphis  {Phorodon  humuli).  I  cannot  believe,  however, 
that  they  had  been  actually  passed,  in  spite  of  the  case  being  reported  by  a  medical  man.— 


PEDICULUS   VESTIMENTI 


533 


each  provided  with  a  pair  of  stigmata.  The  thorax  is  as  broad  as  the 
abdomen.  Eggs  o"6  mm.  in  length;  about  fifty  are  deposited  by  a 
female  head  louse.     The  young  can  propagate  when  eighteen  days  old. 

The  head  louse  lives  especially  in  the  hairy  parts  of  the  head 
of  human  beings  ;  more  rarely  it  is  found  on  other  hairy  parts 
of  the  body.  It  is  spread  over  the  entire  surface  of  the  globe, 
and  was  present  in  America  before  the  arrival  of  Europeans. 
Quite  exceptionally  it  is  said  that  it  bores  itself  deep  into  the 
epidermis  and  can  live  in  ulcers, 

[The  eggs  are  pear-shaped  and  are  attached  to  the  hairs 
near  the  roots  by  means  of  a  clasping  collar.  They  hatch 
in  about  seven  days.  The  young  are  like  the  adults  and 
mature  in  a  month.  Its  general  colour  varies  with  that  of 
its  host.  In  West  Africans  nearly  black,  in  Hindoos  dark  and  smoky,  on  Chinese 
and  Japanese  yellow,  on  Hottentots  orange,  on  South  American  Indians  dark  brown 
(Murray).— F.  V.  T.] 


Fig.  374.— Mouth 
parts  of  Pediculus 
vestimenii.  Enlarged. 
(After  Denny.) 


Fig.  375.— Ovum 
of  the  head  louse. 
70/1. 


Fig.  376.— Head  louse,  male.     15/1, 


Fig.  377. — Pediculus  vesti- 
mentis  Burm.  :  adult  female. 
iS/i. 


Pediculus  vestimenti,  Nitzsch,  1818. 

The  head  in  front  is  somewhat  rounded.  Antennae  longer  than 
in  the  head  louse  ;  2  to  3-5  to  4  mm.  in  length  ;  whitish-grey  ;  the 
abdomen  is  broader  than  the  thorax;  stigmata  as  in  P.  capitis. 
Eggs  07  to  o'9  mm.  in  length  ;  about  seventy  are  deposited. 

P.  vestimenti  lives  on  the  neck,  throat  and  trunk  of  persons,  and  the  clothing  next 
the  body,  in  which  also  the  eggs  are  deposited.  The  louse  of  so-called  pedicular 
disease  {P.  tabescentiuin)  is,  according  to  Landois'  researches,  only  the  usual 
P.  vestijnenti ;  moreover,  many  cases  of  phthiriasis  are  attributable  to  mites 
or  fly  maggots. 

[This  parasite  has  often  been  a  great  pest  amongst  soldiers  during  long 
campaigns,  especially  amongst  the  Russians  during  the  Crimean  War.  Vide  also 
notes  in  Addenda  (p.  615)  under  "  Body,  Head  and  Clothes  Lice." — F.  V.  T.] 


534  THE   ANIMAL   PARASITES   OF   MAN 

Genus.     Phthirius,   Leach. 
Phthlrius  inguinalis,  Redi,  1668. 

Syn.  :  Pedicidiis pubis^  L. 
Male  o*8  to  i*o  mm.  in  length  ;  female  1*12  mm.  in  length  ;  colour 
greyish-white ;  form  subquadrate  ;  the  two  posterior  pairs  of  legs  are 
strong  ;  the  abdomen  has  nine  segments  and  six  pairs  of  stigmata  ; 
and  still  another  pair  of  stigmata  is  situated  between  the  two  anterior 
limbs.  Eggs  pear-shaped,  o'S  to  0*9  mm.  in  length,  0*4  to  o'5  mm. 
in  breadth,  and  are  deposited  in  rows  of  about  ten  on  the  hairs. 


Fig.  378. — Phthirius  inguinalis.  Leach  :  they  are  distinguished  by  the  larger 
tracheal  trunks  originating  from  the  stigmata.     Enlarged. 

Pediculus pubis ^  which  is  found  almost  exclusively  in  the  Caucasian  race,  lives  on 
hairy  parts  of  the  body,  but  hardly  ever  on  the  skin  of  the  head  ;  the  pubic 
region  is  its  favourite  place  of  abode. 

[This  species  reproduces  more  rapidly  than  other  lice,  and  is  communicated 
much  more  freely.  The  eggs  are  often  laid  singly  attached  to  the  hairs  near  their 
apex.     It  is  known  as  the  "  crab  louse."— F.  V.  T.] 

(6)   Rhyncota  HEMIPTERA. 
Family.     Acanthiadae. 

Body  flattened,  antennae  four-jointed,  rostrum  three-jointed,  wings  atrophied. 

[This  family,  the  CimicidcB,  includes  the  bed  bugs  ;  the  proboscis,  which  lies  in  a 
groove,  is  of  three  segments  ;  the  front  wings  are  shown  by  two  small  elytra,  there  is 
no  trace  of  hind  wings.     Two  species  are  known  commonly  to  attack  man.— F.  V.  T.] 

Genus.     Cimex,  Linnaeus. 

Cimex  lectularius,   Linnaeus. 
Syn.  :  Aca?tthia  lectidaria^  Fabricius,  1794. 
It  measures  4  to  5  mm.  in  length,  3  mm.  in  breadth  ;  brownish- 
red  ;    eight  abdominal   segments.     The  female  deposits  fifty  whitish 
eggs  at  a  time  (1*12  mm.  in  length)  three  or  four  times  a  year  ;    the 


CIMEX   LECTULARIUS 


535 


entire  development  up  to  complete  maturity  takes  about  eleven 
months.  [They  will  breed  all  the  year  round,  but  less  so  in  cold 
weather.— F.  V.  T.] 

The  bed  bugs  live  in  the  cracks  and  fissures  of  human  habitations,  under  carpels, 
behind  pictures,  in  furniture,  bedsteads,  etc.  ;  hidden  during  the  day,  they  attack 
persons  at  night  to  suck  their  blood.  The  alkaline  secretion  of  the  salivary  glands 
dropped  into  the  wound  around  the  separate  bites  causes  the  so-called  "  wheals." 

The  bed  bugs  were  known  in  bygone  days  by  the  Greeks  (/capjo)  and  the  Romans 
(cimex).  They  were  first  mentioned  from  Strasburg  in  the  eleventh  century,  and 
in  England  about  1500. 

[This  is  the  common  bed  bug  of  northern  latitudes  and  must  not  be  confused  ;vith 
the  tropical  bed  bug  {C.  rotu7tdatus).  The  bed  bug  can  migrate  from  one  house  to 
another;  this  especially  takes  place  when  a  house  is  uninhabited.  They  escape 
from  windows  and  pass  along  walls,  water-pipes  and  gutters,  and  so  reach  adjoining 
houses.  This  noxious  pest  accompanies  man  wherever  he  goes  ;  ships  and  trains 
become  infested,  especially  the  former. 

[A  characteristic  feature  in  this  animal  is  the  peculiar  odour  it  produces,  like 
many  others  in  the  same  group  of  insects.  This  odour  comes  from  a  clear,  oily 
volatile  liquid  secreted  by  glands  in  various  parts  of  the  body.  Although  he  normal 
food  is  man's  blood,  the  bed  bug  can  subsist  upon  moist  wood,  dust  and  dirt  that 
collects  in  crevices  in  floors,  walls,  furniture,  etc.     The  puncturing  mouth  consists  of 


Fig.  379. 


■Head  of  the  bed  bug  from  the  ventral  surface,    a,  the  rostrum  ; 
b^  the  antenna  ;  and  r,  the  eye.     ^o\\. 


a  fleshy  under  lip,  within  which  lie  four  thread-like  hard  filaments  which  pierce  the 
flesh,  the  blood  being  drawn  up  through  the  beak. 

[The  eggs  are  oval,  white,  with  a  projecting  rim  around  one  end,  with  a  lid  which 
is  pushed  off  when  the  young  hatch  ;  they  are  laid  in  cracks  and  crevices  in  batches 
of  from  twelve  to  fifty.  The  t-gg  stage  lasts  from  seven  to  ten  days.  The  larval 
stage  so  gradually  passes  into  the  adult  that  one  scarcely  notices  the  change ; 
during  its  growth  the  skin  is  cast  five  times,  and  at  the  change  the  little  wing-pads 
are  seen,  showing  that  the  adult  stage  is  reached.  The  young  larva  is  at  first  pale 
yellowish- white.  It  resembles  the  parent,  but  has  no  trace  of  elytra.  Although 
eleven  weeks  is  said  to  be  necessary  for  their  development,  the  stages  may  be  gone 
through  much  more  rapidly  ;  Howard  and  Marlatt  ^  give  seven  weeks  in  some 
instances.  It  seems  pretty  certain  that  these  Cimex  only  take  one  meal  of  blood 
between  each  moult  and  another  preceding  egg  laying. — F.  V.  T.] 


'  "  Household   Insects,' 
1896,  p.  37. 


Howard    and    Marlatt,    Bull.    4    (N.S.),   U.S.    Dept.    Agric, 


536  THE  ANIMAL   PARASITES   OF   MAN 

Cimex  rotundatus,  Signoret,  1852. 

[This  bug  is  common  in  warm  climates;  it  is  an  abundant  insect 
in  India,  and  King  has  found  it  in  the  Sudan,  where  C.  leciularius  is, 
however,  the  common  species.  It  is  usually  known  as  the  tropical 
bed  bug.  Signoret's  bug  can  be  told  from  the  other  common  species 
by  the  shape  of  the  pronotum.  In  C.  rotundatus  it  is  uniformly 
convex,  whilst  in  C.  lectularins  the  lateral  edges  are  flat  and  sometimes 
even  concave.  The  abdomen  of  rotundatus  is  also  rather  more 
elongate. 

[This  species  is  of  considerable  importance,  as  according  to  Patton 
it  may  act  partly  as  the  intermediary  host  of  the  piroplasma  of 
kala-azar. 

[Wenyon  found  at  Bagdad  that  Cimex  sp.  would  take  up  Leishmania 
from  Oriental  sore,  and  that  the  parasite  developed  into  flagellate 
form.  Patton  came  to  the  conclusion  that  the  bed  bug  transmitted 
Oriental  sore  in  Cambay,  India,  but  Wenyon  contests  this  view 
{vide  Journ.  Loud.  School  Trop.  Med.,  191 2,  ii,  pt.  i,  pp.  13-26). 
Franchini  {Bidl.  Soc.  Path,  exot.,  1912,  v.  No.  10,  pp.  817-819) 
was  unable  to  connect  Cimex  with  this  disease.  At  present  nothing 
seems  proved.  Besides  their  possible  connection  with  kala-azar,  it 
has  been  shown  by  Howard  and  Clark  {Journ,  Exp.  Med.,  19 12,  xvi, 
No.  6,  pp.  850-859)  that  they  can  carry  the  virus  of  poliomyelitis. 

[This  bed  bug  was  originally  described  from  the  Island  of  Reunion 
in  1852  by  Signoret.  A  similar  insect  was  described  from  Burma  by 
Fieber,  in  t86i,  as  C.  macrocephalus.  This  is  the  same  as  Signoret's 
species. 

[The  distribution  given  by  Patton^  is  as  follows  :  India,  Burma, 
Assam,  Malay,  Aden,  Islands  of  Mauritius  and  Reunion.  Patton  in 
this  paper  refers  to  an  erroneous  statement  made  in  a  recent  edition 
of  this  book  (the  last  English  edition).  As  I  have  personally  kept 
ledidarius  in  moist  dirt,  wood  and  refuse  for  over  two  years,  the 
statement  as  far  as  I  am  concerned  is  not  erroneous.  Moreover,  since 
his  doubting  this  fact  the  same  experiment  has  been  twice  repeated 
with  the  same  results.  What  they  did  and  do  persist  on  I  cannot 
say.— F.  V.  T.] 

Cimex  columbarius,  Jenyns. 
[This  is  common  in  parts  of  Europe  in  pigeon  nests,  and  also 
amongst  poultry  {vide  Report  Econ.  Zool.  for  year  ending 
September  30,  1913,  pp.  142-144,  Theobald).  It  occurs  in  Britain  on 
the  latter  and  will  attack  man.  I  have  personally  been  badly  bitten 
whilst  collecting  them.     It  is  rounder  and  has  shorter  antennas  than 


'  Indian  Med.  Gaz.,  February,  1907,  xlii,  No.  2. 


REDUVIID^  537 

C.  lectularlus.  Jenyns  also  described  a  more  pubescent  species  from 
swallows  as  C.  hinindinis.  1  have  recently  received  an  account  of  the 
swallow  bug  invading  a  house  in  Kent  and  causing  much  annoyance. 
— F.  V.  T.] 

Cimex  ciliatus,  Eversmann,  1841. 

3*3  mm.  in  length,  yellowish-red,  thickly  covered  with  hair ; 
indigenous  in  Russia  (Kasan). 

[From  a  single  specimen  seen  it  is  evidently  distinct. — F.  V.  T.] 

Family.     Reduviidae. 

Head  long,  narrowed  behind  into  a  neck  ;  eyes  large,  prominent ;  rostrum 
thick  and  curved  ;  antennae  moderately  long,  slender  at  the  tip  ;  legs  long  and 
stiff;  carnivorous. 

Amongst  the  Rediiviidce  one  genus  is  of  particular  importance, 
namely  the  genus  Conorhinus,  which  has  a  long  head  and  the  first 
segment  of  the  beak  very  much  shorter  than  the  second,  and  the 
posterior  tibiae  longer  than   the  femora. 

These  large  bugs  have  a  wide  distribution,  the  Oriental  region, 
North  and  South  America,  and  the  West  Indies,  Madagascar  and 
West  and  Central  Africa. 

These  large  bugs  may  cause  very  nasty  wounds  by  their  bites,  but 
beyond  that  it  has  recently  been  shown  that  one  interposes  in  the  life- 
cycle  of  a  trypanosome,  namely — 

Genus.     Conorhinus,  Lap. 
Conorhinus  meglstus,  Burm. 

This  large  bug  has  recently  been  shown  by  Chagas  to  be  the  agent 
in  the  development  of  the  trypanosome  (7.  cruzii)  which  is  the  cause 
of  the  well-known  disease  in  many  parts  of  Brazil  called  Barheiro 
(Barbier).  This  insect  is  about  i  in.  long,  black,  with  four  red  spots 
on  the  pronotum,  and  six  red  lateral  lines  on  the  abdomen,  black  legs, 
head  and  beak.  The  insect  is  figured  in  a  coloured  plate  (No.  9)  in 
Mem.  hist.  Oswaldo  Cruz,  IQ09,  i,  fasc.  2,  pp.  158-218. 

A  further  account  is  given  by  Neiva.^ 

Conorhinus  sanguisuga,  Lee.   (Blood-sucking  Cone-nose). 

This  bug  is  also  known  as  the  Texas  or  Mexican  bed  bug,  also  as 
the  big  bed  bug.  It  is  particularly  troublesome  in  the  Mississippi 
Valley  in  bedrooms.  The  bite  is  very  severe  and  results  in  more 
pronounced   swelling    and    inflammation    than   that   of    the   Cimex. 

^  Mem.  Inst.  Oswaldo  Cruzy  1910,  2,  fasc.  2,  pp.  206-212. 


538  THE   ANIMAL   PARASITES   OF   MAN 

Normally  this  genus  feeds  upon  the  blood  of  mammals  and  insects. 

Its  fondness  for  human  blood  appears  to  be  quite  a  new  habit,  and 

appears  limited  to  the  mature  insect  only.     It  is  nearly  an  inch  long, 

flat,  head  very  narrow  and  long,  the  rostrum  short  and  thick.      In 

colour  it  is  dark  brown  with  pink  markings.     They  are  fully  winged 

when   adult,  and  they  fly  with  ease,  entering   houses  on  the  wing, 

especially  being  attracted  by  lights  in  windows  ;  they  also  run  swiftly. 

Like  the  bed  bug  they  conceal  themselves  during  the  day  and  come 

out  at  night  and  bite  the  sleeper.     The  effect  of  the  bite  is  very  varied, 

but  as-  a  rule  a  sore,  itching  wound,  accompanied  by  burning  pain 

and  swellings,  which  may  extend  over  a  good  deal  of  the  body,  occur. 

A  specific  poison  is  undoubtedly  injected  into  the  puncture  ;    but  no 

doubt   serious  results  are  also   due  to  the  beak  being  contaminated 

through  the  insects  feeding  upon  foul  carrion.     Mr.  Lembert,  when 

bitten    by   a   Coiiorhimis   sp.    (?)  on  the  Pacific  slope,  exhibited  the 

following   symptoms  :    an    itching   sensation    extending   up    the  leg, 

large    blotches    manifesting   themselves   on    the    upper   part   of    the 

limb   and   extending   up  to  the    hands   and   arms ;    his  lips  swelled 

and   the   itching  and   swelling   extended  over  the  head  ;    there  was 

also  much  nausea.     Similar  results  are  recorded  from  other  regions.^ 

The  eggs  of  the  C.   sanguisuga   are  at   first    white,  then  become 

yellow,  then  pink ;  the  young  hatch  in  twenty  days.     There  are  two 

larval  and  two  pupal  stages,  the  latter  showing  wing-pads.     The  eggs 

are  laid  and  the  young  feed  out  of  doors,  chiefly  upon  insects.     It  is 

particularly  abundant  in  April  and  May  indoors. 

Conorhinus,  sp.  novum  (Monster  Bug). 

Another  species;  acts  in  a  very  similar  way  in  California,  the 
bite  being  very  poisonous. 

Conorhinus   rubrofasciatus,  de  Geer^  (Malay  Bug). 

This  large  bug  attacks  man  in  Malaysia  and  elsewhere.  It  is  recorded 
as  inflicting  "a  very  nasty  sting,  which  is  done  by  the  huge  proboscis." 
Acute  pain  and  inflammation  follow^  in  a  few  minutes.  In  one  case 
the  whole  leg  became  swollen.  This  species  occurs  over  the  whole 
Oriental  region,  in  Madagascar  and  Sierra  Leone.  It  is  dark  brown 
in  colour  with  dusky  yellow  or  brick-red  markings  on  the  pronotum 
and  elytra.  Donovan  suggests  that  it  may  be  connected  with  the 
kala-azar  piroplasma. 


'  "  Household  Insects,"  p.  42. 

2  \_First  Report  Econ.  Zool.y  1903,  p.  130.— F.  V.  T.] 


REDUVIUS  539 

Conorhinus  renggeri,  Herr-Schaff 
(Great  Black  Bug  of  Pampas). 

This  large  black  bug  is  mentioned  by  Darwin/  who  states  as 
follows  :  ^'  At  night  I  experienced  an  attack  (for  it  deserves  no.  less 
a  name)  of  the  benchuca,  a  species  of  Reduvius,  the  great  black 
bug  of  the  Pampas.  It  is  most  disgusting  to  feel  soft  wingless 
insects,  about  an  inch  long,  crawling  over  one's  body.  Before  suck- 
ing they  are  quite  thin,  but  afterwards  they  become  round  and 
bloated  with  blood,  and  in  this  state  are  easily  crushed.  One  which 
I  caught  at  Iquique  (for  they  are  found  in  Chili  and  Peru)  was  very 
empty.  When  placed  on  a  table,  and  though  surrounded  by  people, 
if  a  finger  was  presented  the  bold  insect  would  immediately  protrude 
its  sucker,  make  a  charge  and,  if  allowed,  draw  blood.  No  pain  was 
caused  by  the  wound.  It  was  curious  to  watch  its  body  during  the 
action  of  sucking,  as  in  less  than  ten  minutes  it  changed  from  being 
flat  as  a  wafer  to  a  globular  form.  This  one  feast,  for  which  the 
benchuca  was  indebted  to  one  of  the  officers,  kept  it  fat  during  four 
whole  months,  but  after  the  first  fortnight  it  was  quite  ready  to  have 
another  suck."     Mr.  Kirby'^  also  refers  to  this  species. 

Conorhinus  variegatus  (Variegated  Cone-nose). 

Occurs  in  Florida  in  houses,  and  chases  bugs  (Cimex)  and  flies ; 
not  definitely  known  to  bite  man. 

Conorhinus   nigrovarius. 

This  species  occurs  in  South  America.  It  is  one  of  the  forms 
known  as  bichuque.     Its  bite  makes  a  troublesome  swelling. 

Conorhinus  protractus 

also  attacks  man  in  Utah.^     It  has  been  called  the  '^big  bed  bug." 

Genus.     Reduvius,  etc. 
Reduvius  personatus,   Linne. 

Syn.  :    Reduviics  personatus,  Leconte,   1855' 

European,  but  also  found  in  the  United  States.     The  bite  causes 

mtense  pain.     It  bites  when  caught  or  handled,  but  does  not  seem 

to   do  so  voluntarily.     Swelling  and  irritation  result  which   may  last 

a  week,  and  may  even  cause  death.*     In  1899  it  was  very  abundant 

'  Charles  Darwin,  "A  Naturalist's  Voyage"  (Voyage  of  the  Beagle),  1888,  p.  330. 
'-2  "  Text-book  of  Entomology,"  1885,  p.  205. 

■^  "  The  Big  Bed  Bug  of  the  Far  West,"  Bull.  18  (N.S.),  U.S.  Dept.  Agric,  1898,  p.  loi. 
^  "  Insects  to  which  the  name  '  Kissing-bug '  became  applied  during  the  summer  of  1899/' 
Bull.  22  (N.S.),  U.S.  Dept.  Agric,  1900,  p.  24. 

34 


540 


THE   ANIMAL   PARASITES   OF   MAN 


at  Washington  and  elsewhere;  other  species  occurred,  and  so  no 
definite  opinion  existed  as  to  the  actual  biter,  but  some  people  took 
R.  personatus  actually  biting.  It  was  first  described  as  a  parasite  of 
man  in  America  by  Walsh  and  Riley.^ 

A  popular  name  for  this  bug  is  the  wheel  or  masked  bug— a 
black  insect,  three-fourths  of  an  inch  long.  The  larva  of  this 
hug  is  carnivorous  and  covers  its  body  with  dust  so  as  to  conceal 
itself  from  its  prey.     The  adult  is  active  on  the  wing. 

Coriscus  subcoleoptratus,  Kirby,  1837. 

Syn.  :  Nabicula  subcoleoptrata,  Kirby,  1837;  Nabis  subcoleoptratus, 
Reuter,  1872;  Coriscus  subcoleopij-aius^  Stal,  1873. 

Northern  United  States.  Howard  was  bitten  by  one  between  the 
fingers— the  pain  was  intense,  like  a  needle  prick,  but  the  swelling 
■was  small.     No  other  case  known. 

Rasahus  biguttatus,  Say,   1831. 

Syn.  :  Pirates  biguttatus,  Stal,  1862  ;  Callisphodrus  biguttatus,  Stal,  1866  ; 
Rasahus  biguttatus,  Stal,  1872. 

Common  in  southern  United  States,  and  found  in  Cuba,  Panama 
and  Para,  etc.  Known  as  the  two-spotted  corsair  on  account  of  the 
great  spot  on  the  hemielytra.  Frequently  found  in  houses,  where 
it  chases  the  bed  bug.  It  also  bites  man  frequently.  From 
1869  Walsh  and  Riley  placed  it  amongst  the  parasites  of  man. 
In  the  United  States  Davidson^  is  of  opinion  that  all  cases  attributed 
to  spider  bites  are  due  to  this  insect. 

Melanolestes  morio,  Erichson,  1848  (Non-walker). 

Syn.:  Pirates  morio,  Erichson,   1848  ;   Melanolestes  morio,  Stal,  1866; 
Pirates picipes,  Herrich-Schaffer,  1848  ;  Melanolestes picipes,  Howard,  1900. 

Guiana  and  Mexico  and  eastern  and  southern  United  States. 
Length  20  mm.,  hides  under  stones  and  logs  during  daylight,  and  flies 
at  night.  Attracted  by  lights  into  houses.  Very  abundant  in  1899 
at  Washington.     Howard  cites  cases  where  it  was  proved  to  bite  man. 

Melanolestes  abdominalis,  Herrich-Schaffer,   1848. 
Syn.  :  Pi7'ates  abdominalis,  Herrich-Schafter  ;  Mela7tolestes  abdominalis,  Uhler,  1875. 
Allied  to  the  former ;   some  say  similar,  but  can  be  told  by  the 
shorter  wings  on  the  female.    It  occurs  in  the  same  localities  as  M.  morio. 

*  American  Entomolbgist,  1 869,  i,  pp.  84-88. 

2  R.  Blanchard,  "  Sur  la  Piqiire  de  quelques  Hemipteres,"  Arch,  de  Par.,  1902,  p.  145. 

•'  "So-called  Spider-bites  and  their  Treatment,"  Therap.  Gaz.,  February  19,  1875. 


ARADID.^.,    LYG^ID^  54I 

Phonergates  bicoloripes. 

This  reduvid  attacks  man  in  Africa. 

Family.     Aradidae. 

Broad  and  very  flat  bugs,  with  antennas  of  four  segments  and  the  beak  of  three  ; 
scutellum  short,  no  cuneus  to  elytra  and  the  tarsi  of  two  segments.  They  normally 
live  under  the  bark  of  trees,  etc.,  and  are  found  in  most  parts  of  the  world. 

Dysodius   lunatus,   Fabr.  (Pito  Bug). 

A  large  species  which  is  found  in  South  America,  frequenting 
houses,  and  bites  very  severely. 

The  Ochindundu. 

The  bug  is  described  by  Wellman  {Joiini.  Trap,  Med.,  April  2, 
1906,  p.  97)  as  not  only  feeding  on  ticks,  such  as  Ornitliodorits 
moubata,  but  as  also  attacking  man.  It  is  called  by  the  Angola  Bantus 
Ihe  ochindundu.  It  is  black  in  colour  ;  the  first  two  pairs  of  legs 
are  of  a  bright  red  hue.  It  has  curious  paddle-like  structures  on  the 
front  four  legs,  which  seem  to  be  designed  for  securely  holding  the 
ticks.  It  infects  native  kraals  for  the  sake  of  preying  on  ticks.  The 
natives  also  state  that  it  inflicts  a  bite  which  far  exceeds  in  painfulness 
that  of  the  tick.  They  compare  the  bite  with  that  of  a  poisonous 
snake. 

Family.     Lygaeidae. 

Scutellum  short  ;  antennjE  four-jointed ;  ocelli  present  ;  membranous  part 
of  hemielytra  with  never  more  than  five  nervures.  Nearly  all  vegetable  feeders . 
A  few  are  recorded  here  as  bitinsf  man. 


Lyctocorls   campestris,  Fabricius. 

Syn.  :  Acafithia  campestris,  Fabr.  i^Lyctocoris  doinesticus). 

Rare  in  habitations,  lives  on  human  blood.  Found  by  Blanchard 
in  a  bed  at  an  hotel  at  Liverpool.  The  bite  is  undoubtedly  worse 
than  that  of  Cimex ;  cosmopolitan.  In  colour  it  is  ferruginous, 
shining,  legs  testaceous ;  hemielytra  slightly  shorter  and  narrower 
than  the  abdomen ;  membranous  portion  transparent,  the  apex 
broadly  fuscous.     Length  3*8  to  4*8  mm. 

Rhodinus   prolixus,   Stal,  1859. 
Sometimes    attacks    man,    and    the    bite    is   very   painful.      It   is 
25    mm.  long  and   8    mm.  broad,   and    occurs   in    Colombia.     It^  is 


542 


THE    ANIMAL   PARASITES   OF   MAN 


found   also   in    Cayenne    and    Venezuela.      This   like    other    species 
is  known  in  South  America  as  bichuque  or  benchuca. 

[A  few  other  unimportant  species  are  also  recorded  as  biting  man, 
such  as  Harpactor  cnientas,  in  the  South  of  France ;  Eulyes  amcena, 
from  Borneo  and  Java  ;  Ariliis  carinatiis,  Forster,  from  Brazil. 
The  latter  appears  to  be  the  same  as  the  Acanthia  serrains, 
Fabricius.— F.  V.  T.] 

Order.  Orthoptera. 

[The  only  Orthoptera  recorded  as  doing  actual  harm  to  man  are  certain  wing- 
less locusts  found  in  Africa.  The  cysticercus  stage  of  a  small  tapeworm  found  in 
rats  and  man  has  been  found  in  an  earwig  (Alcock). 

[The  strange  Hemi77ierid(P  found  in  West  Africa,  resembling  wingless  cock- 
roaches, are  parasitic  on  rats  {Criceto/nys).  PhasinidcE^  or  stick  insects,  are  said  to  be 
able  to  eject  a  fluid  which  may  cause  blindness  if  it  comes  in  contact  with  the  eyes. 

Locusts  Injurious  to  Man. 

[A  wingless  locust — Enyaliopsis  durandi^  Luc— is  recorded  by  Wiggins^  as 
injurious  to  man  in  Uganda.  "The  bite  of  this  insect,"  it  is  said,  "gives  rise  to  a 
very  nasty  eruption,  which  may  extend  over  the  whole  body,  with  high  temperature 
and  general  malaise.  The  skin  at  the  site  of  the  bite  sloughs  away,  and  generally 
leaves  a  large  deep  cavity,  which  heals  very  slowly." 

[An  allied  species — E.petersi^  Schaum — emits  a  clear  yellow  fluid,  but  accordmg 
to  Marshall  this  does  no  harm.^  Stannus  writes  that  "  for  some  years  I  have  been 
cognizant  of  the  fact  that  among  the  natives  of  Nyasaland  an  allied  if  not  the 
same  species  is  held  to  cause  skin  lesions  by  the  emission  of  a  fluid  on  the  bare  skin 
surface  of  the  body.  I  have  seen  cases  of  ulcers  on  various  parts  of  the  body,  for 
which  the  '  nantundua  '  was  assigned  as  the  cause."  He  then  describes  the  destruc- 
tion of  the  superficial  layers  of  the  skin  which  he  observed  after  the  yellow  fluid  had 
been  on  the  skin  twelve  hours. — F.  V.  T.] 

Order.  Coleoptera. 

The  larvae  of  beetles,  similarly  to  those  of  some  other  Arthropoda  (myriapods 
and  the  larvce  of  gnats),  have  sometimes  been  observed  in  man  as  purely  accidental 
guests.  In  one  case  or  another,  such  accounts  may  have  originated  through  a 
mistake  of  the  observer.  Thus  English  doctors  report  the  presence  of  the  larvae  of 
Blaps  niortisaga  in  the  stools  of  human  beings,  Sandberg  of  the  larva  of  Agrypnus 
murinus  in  his  ten  year  old  son,  and  Blanchard  mentions  the  larva  of  a  beetle  that 
was  vomited  by  a  child.  All  these  cases,  however,  do  not  represent  actual  parasitism, 
although  there  are  beetles  living  parasitically.^ 

Silvanus  surinamensis,  Linnaeus  (Saw-toothed  Grain  Beetle). 

[Taschenberg  records  this  beetle  as  having  invaded  some  sleep- 
ing apartments  adjoining  a  brewery  where  stores  were  kept,  and 
annoying  the  sleepers  at  night  by  nipping  them  when  in  their  beds. 

1  Bull.  Ent.  Res.,  1910,  i,  pt.  3,  p.  227.  '^  Ibid.,  191 1,  ii,  pt.  2,  p.  180. 

3  [Dr.  Daniels  has  sent  me  a  small  coleopterous  larva  found  in  an  abscess  on  a  man  in 
British  Guiana.— F.  V.  T.] 


DIPTERA  543 

[This  beetle  is  common  in  many  parts  of  the  world  amongst 
groceries,  corn,  meal,  seeds,  dried  fruits,  etc.  It  is  about  ^^  in.  long, 
much  flattened  and  chocolate-brown  in  colour.  The  thorax  has  two 
shallow  grooves  and  bears  six  minute  teeth  on  each  side.  The  jaws 
are  strong,   but  the  bite  cannot  be  very  serious. — F.  V,  T.] 

Order.     Diptera  or  Siphonaptera. 
Aphaniptera  (Fleas). 

Wingless,  the  thoracic  rings  distinct  and  free  ;  antennae  of  three  segments  ;  legs 
very  powerful ;  abdomen  with  nine  segments.  [Ten  segments  are  present,  but  only 
nine  are  visible. — F.  V.  T.]  The  mandibles  transformed  into  serrated  puncturing 
organs,  which  are  situated  in  the  split  sheath  of  the  rostrum ;  the  maxillae  are 
laminated  and  have  palpi,  and  more  or  less  conceal  the  other  parts. 

The  importance  of  fleas  lies  mainly  in  the  fact  that  they  act  as  plague  carriers. 
About  150  species  have  already  been  described.  The  only  ones  of  importance  for 
this  work  are  those  found  on  man  and  those  on  rats  and  mice.  The  two  families 
in  which  these  are  found  are  known  as  Pulicidce  and  Sarcopsyllidce. 

The  eggs  of  fleas  are  laid  on  the  ground,  on  rugs,  etc.,  and  in  birds*  and  rodents' 
nests.  They  hatch  rapidly  in  warm  weather  and  in  warm  climates,  varying  from 
two  to  five  days  ;  in  cold  countries  they  may  take  two  or  three  weeks  to  incubate. 

The  larva  is  a  footless  creature,  pearly  white  in  colour,  the  head  sometimes 
being  darkened,  composed  of  fourteen  segments  including  the  head,  and  although 
apodal  can  move  with  considerable  agility.  It  lives  amongst  dust  and  dirt,  and 
feeds  upon  any  organic  matter  it  can  find.  In  about  two  weeks  it  is  said  to  become 
mature,  and  then  spins  a  cocoon  in  which  pupation  takes  place. 

The  cocoons  of  the  common  human  flea  and  the  fowl  flea  become  covered  with 
dust  and  dirt.  The  period  of  pupal  life  seems  varied,  for  I  have  had  the  fowl  flea 
hatch  out  in  ten  days,  and  others  in  three  weeks  at  the  same  time  of  year. 

The  adults  are  a  blood  suckers  and  cause  considerable  irritation  as  well  as 
acting  as  disease  carriers,  and  in  the  Sarcopsyllidce  the  females  attach  themselve:> 
permanently  to  their  hosts,  embedding  themselves  under  the  skin,  where  they 
become  pregnant.  Some  kinds  harbour  the  cystic  stage  of  tapeworms,  and  the  rat 
trypanosome  passes  certam  stages  in  the  rat  flea.  Most  fleas  have  definite  hosts, 
but  some,  like  the  rat  and  fowl  fleas,  attack  man. 

The  fleas  which  can  carry  the  bacillus  of  plague  are  Xenopsylla  cheopis,  Pulex 
irritans,  Ceratophyllus  fasciatus  and  Hoplopsyllus  anomalus. 

The  two  families,  PulicidcB  and  Sarcopsyllidcr,  can  be  distinguished  as  follows  : 
Thoracic  segments  much  foreshortened,  coxae  and  femora 

of  hind  legs  very  slightly  enlarged      Sarcopsyllidce. 

Thoracic  segments  normal,  coxae  and  femora  of  all  the 

legs  much  enlarged         ...         ...         ...     Pulicidce. 

Family.     Sarcopsyllidae  (Jiggers). 

The  members  of  this  family  are  not  confined  to  one  host. 
Three  genera  are  known  and  tabulate  as  follows  : — 
a.  Hind  coxa  without  a  patch  of  spines  on  the  inside. 

aS.  Hind  femur  simple      \.  Dermatophilus. 

or.  Hind  femur  with  a  large  tooth-like  projection  near 

the  base 2.  Hectopsylla. 

j3.  Hind  coxa  with  a  patch  of  short  spines  on  the  inside        3.  Echidnophaga. 


544 


THE   ANIMAL   PARASITES   OF   MAN 


Genus.     Dermatophilus,  Guerin. 

Dermatophilus    caecata,    Enderl. 
The  eyes  of  the  female  vestigial.     Taken  on  and  behind  the  ears  of  Mus  rattus 
in  Brazil. 

Dermatophilus  penetrans,  L.,  1758  (Jigger,  Chigoe). 

S  y  n . :  Sarcopsylla  petieirans. 

About    I  to   i"2  mm.  in  length;    brown  in  colour.     Eyes  distinct. 

The  males  only  occasionally  visit  man  to  bite ;   the  fertilized  female, 

on  the  other  hand,  bores  into  the   skin  with  her  head,  particularly 

about  the  toes  of  the  host,  and  then  attains  considerable  dimensions. 

The    eggs     develop    on    the 
I       soil    with    a    metamorphosis 
!       similar      to 
common  flea. 


Fig.     380. —  Dermatophilus   fetietraus  :    young 
female.     Highly  magnified.     (After  Moniez.) 


FiCt.    381.  — Dermatophilus    penetrans: 
older  female.    Enlarged.     (After  Moniez.) 


The  sand  flea  (nigua)  particularly  infests  Central  and  South  America,  and, 
in  1873,  was  carried  by  ships  from  Brazil  to  the  West  Coast  of  Africa.  In  a  com- 
paratively short  time  it  has  become  disseminated  throughout  Africa  and  has  also 
appeared  in  Madagascar  ;  recently  also  it  has  been  reported  from  China. 

Besides  attacking  man,  it  also  settles  on  mammals,  for  instance,  on  dogs,  pigs, 
etc.  According  to  Jullien  the  wound  or  little  swelling  caused  by  the  female  has  no 
particular  significance,  as  children  infested  with  ten  or  eleven  sand  fleas  quietly 
proceeded  with  their  ganjes.  It  will  be  understood,  however,  that  the  wound  easily 
affords  the  opportunity  for  the  setting  up  of  inflammation  or  even  septic  processes, 
as  is  the  case  in  any  kind  of  wound. 

[The  jigger  is  also  well  known  in  the  West  Indies.— F.  V.  T.] 

Genus.     Echidnophaga,  Ollifif. 
Four  species  found  on  rats,  etc. 

Echidnophaga  gallinacea,  Westwood  (Chigoe  of  Fowls). 
[This  flea  is  a  native  of  tropical  Asia  and  Africa.     It  lives  on  the 
fowl  chiefly,  attacking  the    neck  and  around  the  eyes.     Specimens 


PULICID^ 


^545 


Pulex, 

Xeiiopsylla. 

Hoplopsylhcs. 

Ctenocephalus. 


were  sent  me  from  Texas,  where  they  not  only  attack  poultry  but  also 
children,  the  latter  somewhat  severely.  It  also  occurs  on  cats,  iand  is 
found  on  rats  in  Africa.  It  has  been  introduced  into  North  America. 
[Three  other  species  are  found  on  rats,  viz.  :  E.  jiiyrmecobii, 
Rothsch.,  from  Australia ;  E.  imirina,  T'lrdh,  irom.  Southern  Europe ; 
and  E.  liopns,  Rothsch,,  India  and  Western  Australia. — F.  V.  T.] 

Family.     Pulicldae  (True  Fleas). 
Rothschild's  classification  is  as  follows  : — 

Section  I. — Club  of  antennas  distinctly  segmented  only  on  the  hind  side. 

A>K  /<?  Genera, 
a.  No  comb  on  head  and  thorax. 

a\  The  internal  incrassation,  which  extends  from  the  inser- 
tion of  the  mid  coxa  into  the  thorax,  joins  the  anterior 

edge  of  the  mesosternite 

dK  This  incrassation  joins  the  upper  edge  of  the  mesosternite 
d.   With  a  comb  on  the  pronotum  only 
.    With  a  comb  on  the  pronotum  and  at  the  lower  edge  of  the 
head       ...         

Section  II.— Club  of  antennae  distinctly  segmented  all  round. 

Key  to  Genera, 
a.  Eye  developed. 

a^.  No  comb  on  head. 

a"-.  Pygidium     not     projecting     backwards  ;     frons     with 

tubercle     ...         ...         

b'.   Pygidium  strongly  convex,  projecting  backwards  ;  frons 
without  tubercle  ... 

b^.  Two  spines  at  angle  of  gense  ...         ...         

h.  Eye  vestigial  or  absent. 
a^.  Abdomen  without  comb. 

a^.  Hind  edge  of  tibiae  with  about  eight  short  and  several 
long  bristles,  which  do  not  form  a  comb. 
a^.  Fifth  segment  in  fore  and  mid  tarsi  with  five,  and  in 

hind  tarsus  with  four  bristles 
b^.   PMfth  segment  in  fore  and  mid  tarsi  with  four,  and  in 
hind  tarsus  with  three  lateral  bristles,  there  being 
an  additional  pair  of  bristles  in  all  the  tarsi  on  the 

ventral  surface  in  between  the  first  pair     

<$>-.   Hind  edge  of  tibias  with  about  twelve  short  and  three 
long  bristles,  the  short  ones  forming  a  kind  of  comb 
b^.   Abdomen  with  at  least  one  comb      


Ceratophyllus. 

Pygiopsylla. 
Chiastopsylla. 


Neopsylla. 


Ctefiophthalmus . 

Ctenopsylla. 
Hystricopsylla. 


Genus.     Pulex,  Linn. 

Pulex  irritans,  L.,  1758. 

Male  2  to  2*5  mm.  in  length,  females  about  4  mm. ;    reddish  or 

dark  brown  ;  head  without  bristles;  thoracic  and  abdominal  rings  of 

bristles  on  the  dorsal  aspect,  and  small  hairs  directed  backwards  at 

he  posterior  margin.      The  barrel-shaped  white  eggs  are  deposited 


54<> 


THE  ANIMAL   PARASITES   OF   MAN 


in  cracks  in  the  boards,  sweepings,  spittoons,  etc. ;  they  produce  legless 
larvae  consisting  of  fourteen  segments,  which,  after  about  eleven  days, 
are  transformed  into  pupae;  after  another  eleven  days  the  flea  emerges. 


,^--v"'''  !   ' 

m^ 

C;; 

^■" 

\. 

"I 

■  I 

Fig.  382. — Piilex  irritans.     14/1. 


Fig.  38^. — Larva  of  flea. 
Enlarged.'  (After  Railliet.) 


Fig.  384. — Pulex  serraticeps.     22/1. 

BonL  Fleas  live  in  human  dwellings  all  over  the  world,  and  periodically  pass  on  to 
persons  to  suck  their  blood.  They  may  deposit  their  eggs  on  very  uncleanly 
individuals,  and  even  undergo  development,  therefore  it  is  possible  to  find  larvae 
and  pupae  on  such  persons. 

The  dog  flea,  Pulex  serraticeps,  is  easily  distinguished  from  the  flea  of  man  by 
the  large  thick  bristles  on  the  posterior  margin  of  the  first  thoracic  ring  (fig-  384). 

Genus.     Xenopsylla,  Glink. 
Xenopsylla  cheopis,  Rothschild. 
This  is  the  common   rat  flea  of  tropical  countries.     Rothschild  ^ 
says  :  "Although  practically  cosmopolitan,  it  cannot  apparently  flourish 
in  temperate  and  cold  cHmates." 


'  Bzill.  Ent.  Res,,  191 1,  i,  pt.  2,  p.  92. 


CTENOCEPHALUS,    HOPLOPSYLLUS,    CERATOPHYLLUS  547 

In  the  male  the  bristles  of  the  flap-like  process  of  the  clasper  all 
slender ;  in  the  female  the  narrow  portion  of  the  receptaculum 
seminis  long.     Originally  discovered  in  Egypt. 

This  is  apparently  the  chief  plague  flea.  The  Indian  Plague 
Committee  have  proved  that  this  flea  is  easily  infected  when  fed  on 
plague  rats,  and  that  the  bacillus  multiplies  rapidly  in  the  flea's 
stomach  and  that  the  fleas  may  remain  infective  for  fifteen  days.  How 
the  flea  infects  man  does  not  apparently  seem  to  have  been  proved,  as 
it  does  not  do  so  through  its  bite,  but  the  excrement  is  highly  infec- 
tive. It  is  probable  that  this  poisoned  faecal  matter  gets  to  the  wound 
caused  by  the  piercing  mouth. 

Xenopsylla  brasillensis,   Baker, 
occurs  on  rats  in  West  Africa  and  has  been  introduced  into  Brazil. 

Genus.     Ctenocephalus,  Kolen. 

Includes  the  cat  and  dog  fleas.  The  dog  flea,  C.  canis,  Duges, 
is  found  on  the  dog  all  over  the  world,  but  especially  in  temperate 
climates.  It  also  occurs  on  rats.  Man  is  often  badly  bitten  by  this 
insect  and  it  overruns  houses.  The  eggs  are  laid  on  rugs,  carpets  and 
dust  and  dirt  and  amongst  dogs'  hair,  but  are  not  fastened  to  it  and 
fall  anywhere.  The  ova  may  hatch  in  about  fifty  hours  and  the  larvae 
live  for  seven  days  and  then  spin  their  cocoons  amongst  dust  and  dirt. 
The  pupal  stage  lasts  about  eight  days. 

The  cat  flea  (C.  fells)  is  widely  distributed  over  the  world,  and 
occurs  on  many  mammals  beyond  the  cat,  and  is  also  found  on  rats. 

Genus.     Hoplopsyllus,  Baker. 

A  genus  found  in  North  America  related  to  Pulex,  but  at  once 
recognized  by  the  prothorax  bearing  a  comb. 

Hoplopsyllus  anomalus,  Baker, 

which  is  found  on  the  ground  squirrel  (Citellus  heecheyi)  in  California, 
an  d  according  to  Rothschild  once  found  on  the  rat,  has  been  proved 
to  carry  the  plague  bacillus  and  to  play  an  important  part  in  plague 
infection  in  California. 

Genus.     Ceratophyllus,  Centis. 

Ceratophyllus  fasciatus,  Bosc. 

This  flea  is  also  found  on  the  rat  in  Europe  and  will  attack  man. 
It   is   a   plague    carrier.     It    has   eighteen   to    twenty   teeth   on   the 

'  •'  Report  United  States  Public  Health,  1909,"  xxiv,  No.  29. 


548  THE   ANIMAL   PARASITES    OF   MAN 

prothoracic  comb  and  no  black  spines  on  the  head.  The  genus 
Ceratophylkis  of  Centis  has  a  pronotal  comb  and  three  ante-pygidial 
ch^ta3  on  each  side.  Two  other  specimens  recorded :  C.  londiniensis, 
Kothsch.,  and  C.  anisiis,  Rothsch.  The  former  on  rats  and  mice  in 
London,  the  hitter  on  Felis  sp.  from  Japan  and  Mus  iiorvegicus  in 
Cahfornia. 

Genus.     Ctenopsylla,  Kolen. 
This   genus    contains   a  very   abundant    rat   and    mouse    species, 
C.  rniisciili,  Duges,  which  is  widely  distributed  over  the  globe. 

Genus.     Hystrichopsylla,  Tasch. 

Large  hairy  fie^s,  with  no  eyes  and  one  or  more  combs  on  the 
abdomen.  In  the  Mediterranean  area  one  species,  H.  tripediiiata, 
Tirah,  is  common  on  rats  and  mice,  and  also  in  the  Azores.  Several 
others  occur  on  rats  and  mice.  For  information  concerning  these  the 
reader  is  referred  to  Rothschild's  papers. 

Pulex  pallipes  is  another  species  found  on  the  rat  and  man. 
Systematic,  Anatomical,  and  Biological  Remarks  on  Mosquitoes. 

Mosquitoes  come  in  the  Nejuatocera^  one  of  the  sub-orders  of  the  Diptera^ 
and  are  divided  into  numerous  families,  of  which,  however,  the  Culicidct  are  of  most 
interest  to  us  here.  Other  families  as  the  Psychodidcc  and  Chironomidce  are  also 
of  considerable  importance,  vide  following  pages.  The  head  is  small,  the  facetted 
eyes  are  placed  laterally,  but  there  are  no  accessory  eyes  (ocelli).  In  front  of  the 
eyes  are  situated  the  comparatively  long  antennas,  the  differences  of  which  strongly 
mark  the  distinction  of  sex.^ 

The  antennas  are  composed  of  fifteen  or  sixteen  segments.  In  the  male  they 
are  covered  with  long  whorl-like  hairs,  while  in  the  female  the  antennal  hairs  are 
short— differences  that  are  perceptible  even  with  the  naked  eye.-  The  proboscis, 
which  is  longer  than  the  antennit,  protrudes  from  the  inferior  aspect  of  the  head  and 
is  composed  of  the  following  parts  (figs.  387  and  388) :  Two  grooved  half  tubes, 
facing  one  another,  of  which  the  upper  one  is  the  upper  lip  (labrum)  and  the  lower 
one  the  lower  lip  (labium),  which  represents  a  pair  of  coalesced  maxillas.  Within 
the  tube  formed  by  the  labrum  and  labium  are  the  mandibles  and  maxillae,  trans- 
formed into  instruments  for  piercing,  and  a  single  puncturing  organ,  the  hypo- 
pharynx.  On  the  right  and  left,  next  to  the  proboscis,  are  placed  the  straight  five- 
jointed  palpi,  the  final  joint  of  which  is  thickened  in  the  male.'  In  biting,  the 
labrum.  which  is  swollen  at  its  free  end,  is  not  introduced  into  the  wound  like  the 

*  [This  is  by  no  means  always  the  case  ;  in  the  genera  Deinocerites,  Wyeomyia,  Limatus, 
Theobald,  and  in  Sabethes,  Robineau  Desvoidy,  they  are  nearly  the  same  in  both  sexes. — 
F.  V.  T.l 

'^  [This  is  not  always  the  case,  vide  previous  note. — F.  V.  T.l 

•''  [This  is  only  so  in  Anophelina  and  in  the  genus  Theobaldinella,  Neveu-Lemaire, 
Grabhamia,  Theobald,  Acartomyia,  Theobald,  elc.  In  true  Culex  and  many  other  genera 
the  male  palpi  are  pointed. — F.  V.  T.] 


REMARKS   ON    MOSQUITOES 


549 


other  mouth  parts,  but  is  bent  backwards.  The  labium  and  hypopharynx  push 
direct  into  the  skin  ;  the  maxillae  and  mandibles,  however,  which  are  needle-like  and 
serrated; at  the  tips,  penetrate  with  a  saw-like  movement.  [The  swollen  free  end  of 
the  labrum  really  means  the  labellce^  two  articulated  pieces,  supposed  by  some  to  be 
the  labial  palpi.  In  most  species  the  mandibles  are  not  serrated  at  their  ends. — 
F.  V.  T.]      The  saliva  is    introduced  into  the    wound  through  the  lumen  of  the 


(«) 


(^) 


Fig.  385. — Head  of  a  male  {a)  and  of  a  female  [b)  Anopheles.    Slightly  enlarged. 

(After  Giles.) 


{a) 


(^) 


Fig.  386.— Head  of  a  male  {a)  and  of  a  female  {b)  Culex.     (After  Giles.) 

hypopharynx,  while  the  blood  is  sucked  up  by  the  mosquito  in  the  groove  of  the 
labium. 

The  three  thoracic  segments  are  soldered  together.  The  central  one  carries  the 
membranous  wings  on  the  sides  of  the  dorsal  surface  ;  the  posterior  somite  carries 
the  small  halteres  (rudimentary  posterior  wings).  There  are  three  pairs  of  long 
slender  legs  on  the  lower  side. 


550 


THE   ANIMAL   PARASITES   OF   MAN 


The  abdomen  has  no  limbs,  is  composed  of  eight  (rarely  nine)  distinct  segments  ; 
the  sexual  and  anal  orifices  are  at  the  posterior  end,  the  stigmata  on  the  sides.  The 
intestinal  canal  (fig.  389)  is  composed  of  three  principal  divisions  ;  the  anterior 
part  reaches  as  far  as  the  front  pair  of  legs,  and  consists  of  the  oesophagus,  which 


md. 


■■ 

y     7  H  -— 

,  -^^ipf  J_^  1 

tr 

PI 

\ 

i\  li  J 

-  4\ \\  i 

Iwli  J 

Wi  ^i 

w^y^ 

\VWl// 

L ,--    u.i. 


ind. 


Fig.  387. — Mouth  parts  of  Anopheles  claviger.^  h.,  hypo- 
pharynx;  md.,  mandible;  mx.,  maxilla;  u.L,  upper  lip;  /./., 
lower  lip  ;  f..,  palpi.     (After  Grassi.) 


7nd. 


Fig.  388. — Anopheles  maailipennis  :  transverse  section  through  the 
proboscis  of  a  female  {a)  and  a  male  {b).  hy.,  hypopharynx,  with  duct 
of  the  salivary  gland;  ?«.,  muscles;  md.,  mandibles;  7?tx.,  maxillae; 
/.,  labium  ;  /./.,  labrum.     (After  Nuttall  and  Shipley.) 

is  provided  with   two   small  lateral  diverticula.      [At  the   commencement   of  the 
(esophagus  are  one  or  more  diverticula,  which  vary  in  size  ;  they  contain  air,  food 

'  [This  should  read  Anopheles  maailipennis,  Meig.  ;  there  was  no  type  of  A.  claviger. 
— F.  V.  T.l 


REMARKS   ON    MOSQUITOES  551 

and  bacteria.— F.  V.  T.]  The  mid  gut  reaches  as  far  as  the  fifth  and  sixth  abdominal 
ring ;  in  front  it  is  thin,  and  has  numerous  small  supra-oesophageal  ganglia ;  the 
posterior  part  is,  however,  more  dilated.  Four  or  five  Malpighian  tubes,  the  excretory 
organs,  discharge  at  the  place  where  the  mid  gut  passes  into  the  terminal  gut. 

The  pair  of  salivary  glands  have  one  common  excretory  duct  leading  into  the 
hypopharynx. 

These  glandular  bodies  are  situated  in  the  thorax  ;  each  consists  of  three  slightly 
serpentine  tubules,  the  dorsal  and  ventral  tubes  being  long,  the  central  one  shorter. 
The  above-named  characteristics  apply  to  both  genera  Culex  and  Anopheles,  but  in 
the  genus  Culex  is  smaller,  Anopheles  larger.  [In  Anopheles  the  ends  of  the  ducts 
in  the  lobules  are  dilated,  whilst  in  most  of  the  genera  the  ducts  are  the  same 
size  all  along.  The  lobules  may  bifurcate,  and  in  Psorophora  there  are  five  lobules. — 
F.  V.  T.]  The  legs  of  the  genus  Culex  are  about  the  same  length  as  the  whole  body ; 
in  Anopheles  they  are  double  that  length.^  In  Anopheles  the  palpi  and  proboscis  are 
of  equal  length;  in  Culex  the  condition  is  different,  according  to  sex.  In  the  male 
the  palpi  are  longer  than  the  proboscis  ;  in  the  female  considerably  shorter  and  the 
number  of  segments  diminished.  The  venation  of  the  wings  exhibits  furthur  points  of 
differentiation,  as  also  their  adornment,  though  this  last  sign  is  not  by  any  means 
always  conclusive  ;  most  species  of  the  genus  Culex  have  unspotted  wings,  whilst 
those  of  Anopheles  are  usually  spotted.     More  important  is  the  fact  that  in  Culex  the 


Fig.  389. — Longitudinal  section  of  an  Anopheles,  showing  alimentary  canal.  In  the  forepart 
of  the  thorax  is  the  salivary  gland  consisting  of  three  tubules  ;  ventrally,  the  suctorial  stomach 
extending  into  the  abdominal  cavity  ;  the  stomach,  and  at  the  posterior  end  of  the  abdomen 
the  Malpighian  vessels.     (After  Grassi.) 

abdomen  is  decorated  with  small  scales,  similar  to  those  on  butterflies,  whereas 
there  are  small  bristles  on  the  abdomen  of  Anopheles.  [This  cannot  be  said  to  be  a 
character  by  which  an  Anopheline  may  be  told  from  a  Culicine,  for  in  such  common 
Anopheline  genera  as  Cellia  and  Neocellia  we  get  plenty  of  scales  on  the  abdomen. — 
F.  V.  T.]  An  experienced  observer  can,  however,  separate  the  two  genera  by  the 
difference  in  size  and  their  manner  of  resting.  When  settled  they  either  touch  the 
resting  place  with  all  the  legs  or  only  with  the  four  anterior  legs.  In  consequence 
of  the  different  length  of  the  legs,  the  body  of  Culexj  approaches  the  resting  place 
more  closely  ;  moreover,  Culex  holds  the  abdomen  parallel  or  at  an  acute  angle  to 
the  resting  surface,  whereas  Anopheles  carries  the  abdomen  directed  upwards  (at  an 
angle  of  about  145°)  and  holds  the  head  down.  Both  genera,  however,  usually  only 
rest  on  the  four  anterior  legs,  and  then,  as  has  long  been  known,  Culex  carries  the 
third  pair  directed  towards  the  dorsum,  while  those  of  Anopheles  hang  down. 

In  regard  to  the  differentiation  of  the  species,  I  must  refer  you  to  the  special 
literature,  and  content  myself  by  observing  that  about  150  species  of  Culex  and 
about  fifty  species  of  Anopheles  have  been  described,  of  which  fifty  about  four  are 

^  [This  is  certainly  not  always  the  case. — F.  V.  T.] 


552 


THE   ANIMAL   PARASITES   OF   MAN 


found  in  Europe.  [The  number  of  known  Anophelines  now  is  more— loo  species— 
of  other  CuhcidiE  over  700.— F.  V.  T.]  According  to  our  present  knowledge  it 
appears  that  the  entire  genus  Anopheles  can  transmit  malaria  to  man  ;  this  observa- 
tion has  been  confirmed  in  Anopheles  clavi'oer,  Fabr.  ;  A.  inacidipennis,  Meig. ; 
A.  bifurcaius,  L. ;  A.  superpictus,  Grassi ;  A.  pseudopictiis,  Gr.,  all  of  which  are 
found  in  Italy,^  Germany,  etc.,  as  well  as  in  the  tropics.  Moreover,  in  A.  costalis^ 
Loew  ;  A.funestus,  Giles  (Africa) ;  A.  quadrimaciilaius.  Say  (North  America),  and 
A.  rossii^  Giles  ;  the  latter  is  perhaps  identical  with  A.  superpictus,  Gr.,  as  well  as 
with  A.  culidfacies  (India).  {Anopheles  maciilipennis  and  A.  claviger  are  the  same. 
Certainly  neither  viaculipenftis  nor  bifurcaius  has  been  found  in  the  tropics. 
Anopheles  quadrimacidatus,   Say,   is  the   same    as   A.  macnlipe7i7tis.     There  is  no 


Fkx.  390. 


-Anopheles  maculipennis,  Meigen.     Enlarged. 
(After  Grassi.) 


evidence  that  all  Anophelines  carry  malaria,  but  there  is  much  to  show  that  certain 
species  only  are  capable  of  so  doing.  A  list  of  known  carriers  is  given  later. — 
F.  V.  T.] 

Everyone  is  aware  that  mosquitoes  swarm  at  sunset  in  fine  weather,  and  then 
seek  out  human  beings  and  other  warm-blooded  animals  to  take  food.  In  this 
regard,  however,  the  sexes  differ,  for  it  is  almost  without  exception  that  the  females 
only  suck  blood,  while  the  males  subsist  on  the  juices  of  plants  (blossoms  or  fruits).^ 

'  Compare  Ficalbi,  E.,  "  Venti  spec,  di  zanzare  {Culicidce)  ital.  .  .,"  Btdl.  Sor,  ent. 
iial.,  1899,  xxxi  ;  abstracted  in  Centralbl.  f.  Bakt.,  Par.  u.  Infektionsk.,  1900,  xxviii,  p.  397. 

■^  Both  males  and  females  may  be  kept  alive  in  captivity  for  a  long  time  if  given  fruits,  or 
even  only  sugar  and  water.     . 


REMARKS   OX    MOSQUITOES 


553 


After  sucking,  and  when  night  has  fallen,  the  mosquitoes  find  a  place  of  refuge,  for 
which  purpose  they  utilize  the  grasses  or  foliage  of  trees  and  bushes,  or  inhabited  or 
uninhabited  rooms  of  houses,  also  cellars,  stables,  verandahs,  etc.,  where  they  also 
pass  the  day. 

[Some  mosquitoes  bite  in  the  daytime— Stegoymia  and  some  Anophelines  ;  some 
bite  right  into  the  night,  as  Cule.x  fatzgans  and  C.  pipiens.—Y .  V.  T.] 

The  period  required  for  digestion  varies  according  to  the  temperature.  It  takes 
two  days  in  summer,  and  may  take  up  to  ten  days  or  more  in  cool  weather.  After 
digestion  is  complete  more  food  is  taken  up,  this  being  necessary  [in  some  species 
only— F.  V.  T.]  for  the  maturing  of  the  sexual  products  in  the  female. 

It  is  still  unknown  under  what  circumstances  copulation  takes  place  ;^  in  any 


t^ 


Fig.    391. — Larva  of  Anopheles  niaculi- 
pennis,  Fabr.     Enlarged.     (After  Grassi.) 


Fig.  392. 


■LTrvaof  Culex.    Enlarged. 
(After  Grassi.) 


case,  sooner  or  later  the  females  are  fecundated,  and  when  the  ova  have  become 
mature,  and  the  season  is  not  too  far  advanced,  they  seek  a  suitable  place  in  which 
to  deposit  them.^  These  are  larger  or  smaller,  permanent  or  temporary,  collections 
of  standing  water,  pools,  puddles,  lakes,  pits,  water  in  rain-water  barrels,  basins,  etc. 
Nevertheless,  certain  kinds  prefer  certain  waters  ;  thus  Anopheles  {claviger)  maculi- 
pennis  and  several  of  the  Culices  seek  stagnant  water  overgrown  with  swamp  vegeta- 
tion and  decomposing  vegetable  matter;  A.  bifurcatus  and  certain  Culices,  clear 
water  with  some  vegetation  (such  as  fountains  and  the  lakes  in  gardens  and  parks) ; 
Ciilex pipiejis  has  a  preference  for  rain-water  barrels,  even  though  the  water  be  dirty 
and  evil-smelling.  [I  have  found  the  larva?  of  Anopheles  bifurcakis  living  in  great 
numbers  in  ponds  and  lakes  completely  overgrown  with  floating  water-weeds,  and 
those  of  Culex pipieiis  in  liquid  manure. 

'  The  act  of  oopulaiion  in  many  species  is  now  known.  The  female  Culex  has  three 
receptaculae  seminalis,  while  ihe  female  Anopheles  has  one  receptaculum  seminis. 

-  It  is  certain  that  the  females  perish  immediately  after  deposidng  the  ova ;  but  this  does 
not  always  hold  good,  as  a  part  of  them  survive  for  a  few  days.  The  males  die  soon  after 
copulation. 


cc^  THE   ANIMAL   PARASITES   OF  MAN 

Sexua/  Organs  of  the  Mosquito.— 1\\^  female  has  a  pair  of  ovaries,  opening  into 
a  single  tube  by  the  ovarian  tubes  ;  into  the  single  tube  opens  a  duct  coming  from 
the  spermatheca;,  and  also  a  mucous  gland.  The  spermathecae  store  up  the  male 
cells.  The  male  organs  consist  of  two  testes  joined  by  ducts  (vasa  deferentia)  to  the 
ejaculatory  duct  formed  by  their  union.  Each  vas  deferens  is  joined  by  a  bhoit  tube 
with  the  sac-like  vesicula  seminalis.— F.  V.  T.] 

There  is  also  a  difference  in  the  manner  in  which  Culex  and  Anopheles  deposit 
their  ova.  Culex  deposits  two  to  three  hundred  eggs  in  compact  heaps  that  float 
on  the  water,  and  in  which  the  eggs  stand  perpendicularly  one  next  the  other  ; 
whereas  Anopheles  jjiaculipenms  deposits  only  three  or  four  up  to  twenty  eggs, 
united  in  groups  that  float  horizontally  on  the  water  ;  the  eggs  of  A.  bifurcatus, 
again,  are  arranged  in  star-like  groups.  The  eggs  are  about  075  mm.  in  length, 
and  assume  a  dark  hue  soon  after  being  laid.  The  development  only  occupies  a 
few  days.     The  young  larvae  grow  rapidly,  changing  their  integument  several  times ; 


Fig.  393. — Pupa  oi  Anopheles  inacuiipennis,  Meig. 
Enlarged.     (After  Grassi.) 

the  larvae  also  differ  in  the  various  genera,  though  they  have  a  general  resemblance 
(figs.  391  and  392). 

The  long  legless  larva  has  a  flattened  head,  a  fairly  broad,  rectangular,  or 
trapeziform  thorax,  on  which  there  are  bristles,  and  an  abdomen  distinctly  seg- 
mented, and  on  the  segments  of  which  there  are  also  lateral  bristles.  The  situation 
of  the  stigmata  marks  the  difference  between  the  two  genera.  Though  in  both 
genera  the  stigmata  are  at  the  posterior  end  and  on  the  dorsal  surface,  they  are  in 
Anopheles  close  to  the  surface  of  the  body  ;  in  Culex,  however,  they  are  on  the  free 
end  of  a  long  tube  (siphon). 

The  position  of  the  larva  in  the  water  also  differs.  The  larva  of  Anopheles  lies 
almost  horizontally  beneath  the  surface  of  the  water,  the  posterior  border  of  the 
penultimate  abdominal  segment,  upon  which  the  stigmata  are  situated,  being  on  the 
surface ;  whereas  the  larva  of  Culex  hangs  head  downwards  perpendicularly  in  the 
water,  the  point  of  the  siphon  only  touching  the  surface. 

In  about  a  fortnight  the  larva  is  fully  grown  and  becomes  a  pupa.     The  pupa 


CULICID^,   OR   MOSQUITOES  555 

(fig"'  393)5  which  moves  in  jerky  movements,  remains  in  the  water,  but  partakes  of 
no  food.  In  shape  it  somewhat  resembles  a  tadpole,  that  is  to  say,  it  consists  of  a 
bulky  anterior  portion,  on  the  surface  of  which  the  head,  with  its  appendages,  is 
recognizable,  and  a  more  slender  segmented  abdomen.  Above,  on  the  thorax,  there 
are  two  small  trumpet-shaped  breathing  tubes  for  the  conveyance  of  air  to  the 
tracheal  system.  After  three  or  four  days  the  perfect  mosquito  hatches  out,  remains 
a  short  time  on  the  surface  of  the  water  until  its  chitinous  integument  is  hardened, 
and  then  flies  away. 

The  females  that  are  fertilized  in  the  autumn  hibernate  in  sheltered  spots  in  the 
open  air,  or  in  houses,  cellars,  under  stairs,  in  stables,  barns,  etc.,  and  are  the 
progenitors  of  the  first  generation  of  the  following  year. 

In  accordance  with  the  climate  of  a  country,  or  the  kind  of  weather  of  a  year, 
the  conditions  in  regard  to  the  manner  of  life  and  the  duration  of  the  development 
of  the  mosquito  vary.  At  all  events,  the  life-history  of  the  mosquito  elucidates 
many  points  relating  to  malaria  which  were  hitherto  not  understood. 

[The  length  of  the  egg,  larval  and  pupal  life  varies  so  much  that  it  is  not 
possible  to  give  an  account  of  any  value  here.  Frequently  the  eggs  may  incubate 
in  two  days,  whilst  I  have  had  StegoiJiyia  fasciata  eggs  from  Cuba  that  have  hatched 
out  under  abnormal  circumstances  more  than  two  months  after  they  were  laid 
("  Mono.  Culicid.,"  iii,  p.  6).  Some  larvae,  as  Aftopheles  bifurcatus,  live  for 
months  during  the  winter.  Some  mosquitoes  therefore  hibernate  as  larvae.  The 
larvae  and  pup^e  of  the  different  genera  present  very  marked  characters,  mainly  in 
regard  to  the  structure  of  the  siphons.  Specific  differences  may  be  found  in  the 
frontal  hairs  of  Anopheline  larvae  and  in  the  number  and  arrangement  of  a  group 
of  spines  at  the  base  of  the  siphon  in  Culicines. — F.  V.  T.] 


Gulicidse  or  Mosquitoes. 

The  importance  of  these  insects  to  man  is  very  great.  They  not  only  produce 
painful  bites,  which  may  become  inflamed  and  give  rise  to  a  considerable  amount 
of  cedema,  but  they  are  more  important  on  account  of  the  part  they  play  in  the 
distribution  of  various  diseases.  CidicidcE  may  not  only  carry  disease  germs,  but 
act  as  intermediate  hosts  for  certain  parasites,  such  as  some  of  the  Anophelina  for 
malarial  parasites,  Culex  for  Filariae,  and  Stegomyia  for  yellow  fever,  etc.  ;  the  last- 
named  is  in  any  case  the  distributor  of  that  fatal  disease.  It  is  therefore  very 
necessary  to  know  the  life-history,  habits  and  characters  of  these  pests. 

Mosquitoes  exist  in  almost  all  parts  of  the  world  from  the  Arctic  circle  to  the 
tropics  ;  temperate  regions  suffer  from  them  less  than  the  two  extremes,  but  even 
there  they  form  not  only  a  source  of  great  annoyance  but  of  danger  as  malaria  and 
possibly  now  and  again  yellow  fever  carriers.  A  few  years  ago  comparatively  few 
species  were  known,  now  some  800  odd  have  been  described.  Their  number  will 
probably  not  stop  far  short  of  1,000,  in  spite  of  the  fact  that  many  have  been 
described  under  different  names,  yet  really  the  same  species.  Some  are  purely 
domestic,  others  entirely  sylvan  ;  the  former,  as  we  might  expect,  often  have  a  very 
wide  distribution,  having  been  taken  from  place  to  place  in  boats  and  trains.  The 
more  rapid  transport  becomes,  the  greater  becomes  the  possibility  of  this  wide 
distribution  of  many  species  increasing,  and  the  spread  of  other  species  from  their 
natural  home  to  foreign  parts  by  sea  and  then  by  trains  further  inland. 

All  CulicidcE  are  aquatic  in  their  larval  and  pupal  stages.  Almost  all  small 
collections  of  water,  both  natural  and  artificial,  may  form  breeding  grounds  for 
these   pests.      Some  even   breed  in  pitcher  plants  and  many  in   bromelias.     The 

35 


556 


'HE   ANIMAL   PARASITES   OF   MAN 


YiG.  3^^— Heads  of  Culex  and  Anopheles  :    (i)  Culex  trale  ;  (2)  Culex  fema]( 
(3)  Anopheles  male  ;  (4)  Anopheles  female.     (After  Daniels.) 


CULICID^    OR  MOSQUITOES 


557 


favourite  resorts  for  the  larvae  of  Anophelina  are  small  natural  collections  of  water, 
such  as  puddles,  ditches  and  small  pools  around  swamps;  certain  species  (A.  viaculi- 
pennzs,  etc.)  live  in  rain  barrels  as  well.  They  may  also  occur  in  the  sluggish  water 
at  the  edges  of  rivers  or  even  in  mid  river,  where  the  flow  is  checked  by  masses 
of  water  weeds  {Myzomyia  funesta,  etc.).  The  Stegomyias  prefer  artificial  collections 
of  water,  but  also  occur  in  natural  pools.  The  yellow  fever  species  {S.  fasciaid) 
prefers  small  collections,  such  as  in  barrels,  pots,  jars,  etc.  Culex  occur  in  all 
manner  of  places — rain  barrels,  tanks,  cisterns,  ponds  and  ditches.  Some  of  the 
South  American  species  of  Culex,  Wyeomyia,  Joblotia,  etc.,  breed  in  the  collections 
of  water  at  the  base  of  bromelia  leaves.^  Very  few  Culicid  larvae  live  in  salt  water 
except  in  Australia,  where  Dr.  Bancroft  has  found  them  in  salt  water  of  specific 
gravity  1*040  {Mucidus  alternants  and  Culex  ati7iulirostris).  Other  salt  water 
mosquitoes  are  known  in  America.  The  food  of  the  larvae  is  very  varied  ;  the 
majority  appear  to   feed  upon   confervae,   small  Crustacea  and  insects;    some  are 


Fig.  395. — a,  eggs  of  Culex  ;  <5'  b^,  eggs  of  Anopheles  ;  r,  eg^ 
^,  ^goi  Tseniorhynchus  ;  e,  egg  of  Psorophora 


,  egg  of  Stegomyia  ; 


cannibals,  readily  devouring  others  of  their  own  kind.  The  larger  larvae  of  Mega- 
rhinus,  Psorophora,  Toxorhynchites  and  Mucidus  are  extremely  ravenous  and  devour 
one  another. 

There  are  two  main  types  of  larvae,  the  Anopheline  and  Culicine  ;  in  the  former 
there  is  no  respiratory  siphon,  in  the  latter  the  siphon  is  long  or  moderately  long. 
The  head  offers  certain  marked  peculiarities  which  are  of  specific  value  ;  this 
especially  applies  to  the  Anophelina^  in  which  the  frontal  hairs  are  of  great  service 
in  distinguishing  the  larvae,^  whdst  in  Culex  ihe  number  and  position  of  the  spines 
at  the  base  of  and  on  the  siphon  are  characteristic.  The  position  assumed  by  the 
larvae  in  the  water  also  varies  in  the  different  groups  ;  most  of  the  Anophelines  lie 
horizontally,  most  of  the  Culicina  and  ALdeomyina  hang   head  downwards.     The 


1  "  Wald  Mosquitoes  and  Wald  Malaria,"  Dr.  Lutz,  Centralbl.  f.  Bakt.,  Par.,  21. 
Infektionsk.,  i  Abt.  Grig.,  xxxiii,  No.  4. 

2  Informaii  n  sent  me  by  Dr.  Grabharn  shows  this  statement  to  be  not  quite  correct,  as  the 
frontal  hairs  may  vary  in  different  stages  of  the  same  larva.  This  he  has  shown  in  Cellia 
albipes,  Theob.,  and  I  have  noticed  it  in  a  Nyssorhynchus  from  Africa. 


558 


THE   ANIMAL   PARASITES   OF   MAN 


pupee  also  vary,  but  not  to  the  same  extent;  the  chief  differences  to  be  noticed  are 
in  the  form  of  the  two  respiratory  trumpets. 

The  eggs,  which  may  be  laid  separately  {Ajtopheles  macuhpennis^  Stegoviyia 


Proboscis 


Antennae 


1  Wing  scaies"! 


Proboscis 

Palp 

Antennae. 

Basal  lobes  ofantennqe 

Yrorts 

Vertex 

Eyes 

Oaipul— 
Nope 

Fig.  396. — Diagram  showing  the  structure  of  a  typical  mosquito.     (Theobald.) 


5^'?t:arsci 


fasciaia,  Joblotia  nivipes,  etc.),  or  in  rafts  {Ctdex  pipiens^  C.fatigans)  or  in  chains 
{Pseudotceniorhynchus  fasciolatus),  present  a  great  variety  of  forms.  The  most 
peculiar  are  shown  in  fig.  395  (Ttisniorhynchus,  Culex,  Stegomyia,  Anopheles, 
Psorophora). 


CULICID^   OR   MOSQUITOES 


559 


As  in  all  insects,  they  differ  very  materially  in  each  species  of  one  genus.  Those 
best  known  are  the  Anopheline  eggs. 

The  eggs  always  float  on  the  surface  of  the  water;  immersion  soon  destroys 
them,  but  many  may  occur  in  mud  and  can  resist  desiccation. 


Fig.  397. — Types  of  scales,  a  to  k  ;  head  and  scutellar  ornamentation,  I  to  5  ;  forms  of 
clypeus,  6.  (Theobald.)  i,  head  and  scutellum  of  Stegom)ia,  etc.;  2,  of  Culex  and 
Mansonia  ;  3,  of  Howardina,  yEdes,  etc.  ;  4,  of  Megarhinus  and  Toxorhynchites,  etc.  ; 
5,  of  Cellia  and  some  other  Anophelines ;  6,  a',  clypeus  of  Culex;  d',  of  Stegomyia ; 
<:',  of  Joblotia. 


C/tarac/ers  of  Adul^  Cuh'cidcF.— The  ch\e(  ch2iY2iC\.eYS  by  which  true  mosquitoes, 
or  Ciilicidce^  are  known  are  the  following  : — 

(i)  Wings  always  with  the  veins  covered  with  scales;    the  longitudinal  veins, 


56o 


THE   ANIMAL   PARASITES   OF  MAN 


usually  six  in  number  (in  one  genus  seven) ;  the  costal  vein  carried  round  the  border 
of  the  wing. 

(2)  Head,  thorax  and  abdomen  usually,  but  not  always  (Anopheles,  etc.),  covered 
with  scales. 

(3)  Mouth  parts  formed  into  a  long  piercing  proboscis. 

As  a  rule  the  males  may  be  told  from  the  females  "by  their  antennae  being 
plumose,  whilst  in  the  females  they  are  pilose  {vide  fig.  394),  but  this  does  not 
invariably  hold  good,  for  in  Deinocerites,  Theobald,  and  Sabethes,  Desvoidy,  and 
others,  they  are  pilose  in  both  sexes.  The  labial  palpi  are  very  variable  in  regard 
to  their  form  and  the  number  of  segments  ;  in  the  Aiiophelina  they  are  long  in  both 
sexes,  as  long  or  nearly  so  as  the  proboscis,  more  or  less  clubbed  in  the  males ;  in 
Culicma^  Joblotiiia  and  Heptaphlebomyia^  they  are  long  in  the  males,  short  in  the 
females  ;  in  A£deo?nyina,  short  in  both  sexes. 

Scales. — The  most  important  structural  peculiarities  in  Culicidce  are  the  scales, 
which  form  the  chief  and  most  readily  observed  characters  for  separating  genera 
and  species.     The  importance  of  scale  structure  has  been  recently  ignored  by  some 


Fig.  '^98. — Neuration  of  Wing.  Explanation  of  JVing,  Veins  and  Cells. — A,  costal  cell ; 
B,  sub-costal  cell  ;  C,  marginal  cell;  D,  first  sub-marginal  cell  (=  first  fork  cell)  ;  E,  second 
sub-marginal  cell ;  F,  first  posterior  cell ;  G,  second  posterior  cell  (=  second  fork  cell)  ;  H,  first 
basal  cell  ;  I,  second  basal  cell;  J,  third  posterior  cell;  K,  anal  cell;  L,  anxiliary  cell; 
M,  spurious  cell  ;  c,  costal  vein  ;  1st — 6tk,  first  to  sixth  longitudinal  veins  ;  a,  a'  and  a',  incrassa- 
tions  (a' called  by  Austen  the  sixth  vein,  a"  the  eighth  vein);  y,  supernumerary  cross  vein; 
z,  mid  cross  vein  ;  /,  posterior  cross  vein  ;  s.c.^  sub-costal.      (Theobald.) 


workers,  who  are  probably  right  academically,  but  as  a  means  of  separating  groups^ 
and  so  more  easily  running  down  a  species,  the  practical  man  is  strongly  advised  to 
follow  this  method.  As  to  what  a  genus  is,  is  purely  a  matter  of  personal  opinion. 
If  one  examines  any  recent  standard  work  on  entomology  one  will  find  a  species 
being  placed  in  varied  genera  by  the  varied  authorities. 

The  head,  thorax,  abdomen  and  wings  are  in  nearly  all  cases  clothed  with 
squamce  of  varied  form,  of  which  the  following  are  the  main  types  (fig.  397)  : — 

(i)  Flat,  spade-shaped  scales  (a). 

(2)  Narrow  curved  scales  (e). 

(3)  Hair-like  curved  scales  (d). 

(4)  Spindle-shaped  scales  (/). 

(5)  Small  spindle-shaped  scales  (£■). 

(6)  Upright  forked  scales  {/i)  and  (i). 

(7)  Twisted  upright  scales  (J). 

(8)  Inflated  or  pyriform  scales  (/'). 


THE   CLASSIFICATION    OF   CULICID^ 


561 


(9)  Mansonia  scales  (d). 

(10)  Small  broad  asymmetrical  scales  (c). 

Various  other  varieties  are  found  on  the  wings,  such  as  : — 
(i)  Narrow  linear  lateral  scales. 

(2)  Narrow  lanceolate  scales. 

(3)  Broad  lanceolate  scales. 

(4)  Elongated,  broad,  truncated  scales  (=  Pseudotaeniorhynchus-like  scales). 

(5)  Pyriform  scales. 

(6)  Asymmetrical  broad  or  Taeniorhynchus  scales. 

(7)  Flat  spade-like  scales.^ 

The  wings  have  a  series  of  scales  along  the  middle  line  of  the  veins,  and  also 
lateral  scales  to  all  or  nearly  all  the  veins.  The  wing  is  also  fringed  by  a  series  of 
scales  (fig.  396),  which,  however,  are  of  little  systematic  importance  ;  the  so-called 
"  border  scales  "  (b.s.)  vary,  however,  to  some  extent,  and  are  useful  characters  in 
separating  some  of  the  Taeniorhynchus. 

The  Classification  of  Culicidcv. 

Section  A. — Proboscis  formed  for  piercing  ;  metanotum  nude. 
Scutellum  simple. 
I.    Wings  with  six-scaled  longitudinal  veins. 
A.   Palpi  long  in  the  male. 

a.  Palpi  long  in  both  sexes,  clavate  in  S  Aftophelina, 
\.  First  submarginal  cell  as  long  or  longer  than  the  second  posterior  cell. 
Anlennal  segments  without  dense  lateral  scale  tufts. 
/Wing    scales 
lanceolate     ... 
Wing   scales 
mostly    long 
and  narrow  ... 
Wing   scales    as 
above,     but 
f  o  u  r  t  h     long 
vein  near  base 
of    third    and 
outstanding 
scales  on  pro- 
thoracic  lobe 
Wing  scales 
partly   large 
and  inflated 


Thorax  and 
abdomen 
with  hair- 
like scales 


Prothoracic 
lobes  sim- 
ple;  no' 
flat  head 
scales 


Anopheles^  Meigen. 


Myzomyia^  Blanchard. 


Neomyzomyia,  Theobald. 


I  ^  \ 

I  Prothoracic     lobes      mammil- 
'      lated  ;  some  flat  head  scales. 
.     Basal  lobe  of  $  genitalia  of 
\    two  segments 
Prothoracic    lobes    with    dense    outstanding 

scales  ... 
Thorax   with    some    narrow  curved    scales ; 

abdomen  hairy 
Wing  scales  small   and    lanceolate.      Wing 
scales  broad  and  lanceolate 


Cycloleppteron^  Theobald. 

Stethoinyia^  Theobald. 
Feltinella,  Theobald. 
PyretopJwrus^  Blanchard. 
Myzorhynchella^  Theobald. 


Heart-shaped  scales  occur  on  the  wings  of  Etiorlepiiomyia. 


562 


THE  ANIMAL   PARASITES   OF   MAX 


II, 


Thorax  with  hair-Hke  curved  scales,  some 
narrow  curved  ones  in  front  ;  abdomen 
with  apical  lateral  scale  tufts,  scaly  venter  ; 
no  ventral  tuft 

Thorax  with  hair-like  curved  scales  ;  abdo- 
minal scales  on  venter  only,  with  a  distinct 
ventral  apical  tuft      

Much  as  above,  but  abdomen  with  long 
spine-like  dense  lateral  tufts  

Thorax  with  very  long  hair-like  curved  scales ; 
abdomen  pilose,  except  last  two  segments 
which  are  scaly  ;  dense  scale  tufts  on 
third  femora  ;  wings  with  broadish,  blunt, 

lanceolate  scales       

/Abdominal  scales  as  lateral 
dorsal  patches  of  small  flat 
scales ;  thoracic  scales  nar- 

Thorax  and  row  and  curved,  or  spindle- 
abdomen    {      shaped 

with  scales    Abdomen    nearly    completely 
covered  with  irregular  scales 
and  with  lateral  tufts 
^  No  lateral  scale  tufts  ... 

Thoracic  scales  ha[r-like  except  a  few  narrow 
curved  ones  in  front ;  abdominal  scales 
long,"  broad  and  irregular 

Thorax  with  hair-like  curved  scales  and 
some  broad  straight  scales,  others  spatu- 
late  on  sides.  Abdomen  covered  with 
fine  hairs  except  last  three  segments, 
which  are  scaly.  Tufts  of  scales  on  hind 
femora.     Wing  scales  lanceolate 

Antennal  segments  with  many  dense  scaly 
tufts 

Antennas  with  outstanding  scales  on  second 
segnient,  more  appressed  ones  on  the  first. 
At  least  one  segment  of  abdomen  with 
long  flat  more  or  less  spatulate  scales      ... 

First  submarginal  cell  very  small      

With  a  distinct  cylindrical  tubercle  project- 
ing obliquely  from  the  prothoracic  region 

Scutellum  trilobed. 


A rribalzagia^  Theobald. 

My2'.orhynchus^  Blanchard. 
Chrystia,  Theobald. 

Lophosceloniyia,  Theobald. 

Nysso?'hynchus,  Blanchard. 


Cellia^  Theobald. 
Neocellia,  Theobald. 


Kerieszia.  Theobald. 


Mano^uinhosia^  Cruz. 
Chagasia,  Cruz. 


Calver/hia,  Ludlow. 
Bironella,  Theobald. 

Dactyloinyia,     Newstead     and 
Carter.^ 


'  T"^e  following  genera  of  Anophelites  have  been  founded  by  James*  :  — 
(i)   Abdomen  with  hairs  but  no  scales.     Thorax  with  dorsum  with  long  narrow  curved  scales, 
which  form  on  the  anterior  promontory  a  thick  bunch  projecting  over  the  neck.     Protho- 
racic lobes  with  a  tuft  of  rather  broad  true  scales,  upright  forked  scales  of  head  of  usual 
broad  expanding   type:   Patagiamyia,  James.       Includes    Gigas,    Giles,   and    Lindesayi, 
Giles.     Both  seem  to  me  typical  Anoplieles. 
(2)  Abdomen  as  above.    Thorax  very  similar.    Prothoracic  lobes  with  hairs,  no  scales.    Upright 
*  Records  of  Indian  Museum^  1910,  iv,  No.  5,  p.  q8. 


THE   CLASSIFICATION   OF   CULICIDJi: 


563 


First  submarginal  cell  much  smaller  than 
the  second  posterior  cell  ;  proboscis  long 
and  bent         

Palpi  long  in  both  sexes  

Last  segment  of  ^  palpi  blunt.  Last  seg- 
ment of  (^  palpi  long  and  pointed 

Palpi  short  in  the  female 

First  submarginal  cell  longer  than  the  second 
posterior  cell 

Legs  more  or  less  densely  scaly  ;  head  not 
entirely  clothed  with  flat  scales  ;  all  the 
legs  densely  scaly. 

Wings  with  large  pyriform  scales      

Wings  with  narrow  scales 

Hmd  legs  only  densely  scaled 

Head  entirely  clothed  with  flat  scales.  . 

Legs  uniformly  scaled  with  flat  scales. 

Head  and  scutellar  scales  all  flat  and  broad. 

Palpi  of  2  short,  of  ^  thickened  apically 
and  tufted      

Palpi  of  2  longer  than  in  Stegomyia  and  in 
(^  long  and  thin,  acuminate,  simple 

Head  scales  mostly  flat,  but  a  median  line 
of  narrow  curved  ones  ;  scutellar  scales 
flat  on  mid  lobe,  narrow  curved  on  lateral 
lobes  and  palpi  longer  than  proboscis 

Head  scales  mostly  flat,  irregular,  narrow 
curved  ones  behind  ;  mid  lobe  scutellum 
with  flat  scales,  lateral  with  narrow  curved  ; 
^  palpi  shorter  than  proboscis 

Head  scales  mostly  flat,  but  a  few  narrow 
curved  ones  in  middle  in  front ;  scutellar 
scales  all  flat 

Head  scales  all  flat  ;  scutellar  scales  all 
narrow  curved 


Megarhiniiice. 

Megarhinus^  Rob.  Desvoidy. 

Ankylorhynchiis^  Lutz. 
Toxorhyjichites^  Theobald. 

Culiciftce. 


Mucidtis,  Theobald. 
Psorophora^  Rob.  Desvoidy. 
Ja?tthi?iosoma,  Arribalzaga. 


Stegomyia^  Theobald. 
Desvoidea^  Blanchard. 

Macleayia^  Theobald. 

Catageiojnyia,  Theobald. 

Scuto?nyia,  Theobald. 
Skusea.  Theobald. 


forked  scales  of  head  rod-shaped:  Neostelhopheles,  James.  Includes  Atkenii,  James; 
Immaculatus,  Theobald  ;  Culiciformis,  James  and  Liston.  These  seem  to  me  to  be  true 
Anopheles. 

■(3)  Abdomen  with  hairs  and  scales  on  dorsum  of  each  segment ;  ventrally  there  are  six  scaly 
tufts  on  the  apices  of  six  segments.  Thorax  with  scales  and  a  tuft  of  outstanding  ones 
on  prothoracic  lobes  :  Christophersia,  James.  Type  Halli,  James.  Very  close  to  if  rot 
identical  wiih  CelJia. 

{4)  Head  with  narrow  curved  scales  lying  rather  flat  upon  head  and  flat  lateral  scales,  upright 
forked  ones  behind.  Central  lobe  of  scutellum  with  tuft  of  narrow  curved  scales,  lateral 
lobes  with  large  flat  oval  scales  ;  male  palpi  longer  than  proboscis,  two  large  apical  seg- 
ments with  long  projecting  hairs  :  Leslieomyia,  Christophers.  Type  Leslieomyia  tcEuio- 
rhynchoides,  Christophers,  from  Amritsar,  India. 

-(5)  Abdomen  with  first  six  or  seven  segments  with  hairs  only,  eighth  and  seventh  (?)  with 
scales,  also  genital  processes.  Thorax  with  hairs  and  narrow  curved  scales  sharp  pointed, 
blunt-ended  broad  scales  on  each  side  of  anterior  third.  No  tufts  of  scales  on  prothoracic 
lobes.  Head  usual  type  of  upright  forked  scales  :  Nystomyzomyia,  James.  Type  Rossii, 
Giles. 


5^4 


THE   ANIMAL   PARASITES   OF    MAN 


Head  with  flat  scales,  except  a  small  median 
area   of    narrow   curved   ones ;     scutellar 
scales  all  narrow  curved    ... 
Head  with  all  flat  scales  except  a  thin  line 
of  narrow  curved  ones  behind  ;   scutellar 
scales  all  narrow  curved     ... 
Head  with  small   flat   scales  over  most   of 
surface,  with  median  line  and  line  around 
eyes    of    narrow   curved    ones ;     scutellar 
scales  bluntly  spindle  or  club-shaped    ... 
Head   and   scutellar  scales   narrow  curved. 
Wing  scales  long,  narrowly  lanceolate,  col- 
lected   in    spots  ;    palpi    clubbed    in    S  ; 

five-jointed  and  rather  long  in  $ 

Wing  scales  (lateral)  long  and  narrow,  and 
$  palpi  three-jointed,  ^  not  clubbed  and 

hairy 

Wing  scales  at  apex  of  veins  dense  and 
rather  broad,  femora  swollen  ;  small  dark 

species  ...         

Wings  with  short,  thick,  median  scales  and 
short,  broadish  lateral  ones  on  some  of 
the  veins  ;  scales  mottled  ;  fork-cells  rather 

short 

Wings  with  dense,  broadish,  elongated,  trun- 
cated scales   ... 
Wings    with     broad,     short,    asymmetrical 

scales 

Head  covered  with  rather  broad,  flat,  spindle- 
shaped  scales  ;    scutellum  with  small  flat 

scales  to  mid  lobe 

Head  clothed  with  flat,  irregularly  disposed 
scales  all    over,  with    patches    of  narrow 

curved  ones  ;   ^  palpi  clubbed 

Abdomen  with  projecting  flat  lateral  scales 
with    deeply   dentate   apices ;    wings    not 

ornamented    ...         ...         ...         

Wings  ornamented  ;  scutellum  with  flat  and 

narrow  curved  scales  

7.    Palpi  short  in  ^  and  $  

Wings  unornamented. 

Antennae  pilose  in   ^  and   ?  ;  second  joint 

very  long 
Antennas  plumose  in  the  <^. 
Head  clothed  with  narrow  curved  and  flat 
scales. 

Mid-lobe  of  scutellum  with  six  border- 
bristles. 

Scutellum  with  narrow  curved  scales. 

Palpi  in  ?  four-jointed,  in  ^  two-jointed  ... 

Mid-lobe  of  scutellum  with  four  border 
bristles. 


Ho7uardt?ia,  Theobald. 
Danielsia^  Theobald. 

Hulecoctoinyia^  Theobald. 

Theobaldia,  Neveu-Lemaire. 
Ciilex^  Linnaeus. 
Melanoconio?!^  Theobald. 


Grabhamia^  Theobald. 

Pseiidotceniorhy7ichus^ 

Theobald. 
Tceniorhynch  us^  A r  r i  balzaga. 


Giles ia,  Theobald. 


Acarlowyia^  Theobald. 


Lasioco7iops^  Theobald. 

Finlaya,  Theobald. 
A^deoiiiyi7ia. 


Deinocerifes,  T'heobald. 


JLdes^  Meigen. 


HE   CLASSIFICATION   OF   CULICID^ 


565 


Scutelliim  with  flat  scales. 

Head  clothed  with  flat  scales  only. 

Fork-cells  normal  length. 

Mid-lobe  of  scutellum  with  four  border- 
bristles. 

Palpi  of   $  two-jointed  

Palpi  of   ?  five-jointed,  metallic 

Fork  cells  very  small  or  small. 

Scutellar  scales  flat. 

First  SLibmarginal  cell  longer  than  the 
second  posterior  cell;  no  flat  scales  on 
mesothorax    ... 

First  submarginal  cell  smaller  than  the 
second  posterior  cell  ;  flat  scales  on  meso- 
thorax... 

Scutellar  scales  narrow  curved. 

First  submarginal  cell  as  in  Uranotaenia  ... 

Wings  ornamented  with  Mansonia-like  scales 


VeraUi7ia^  Theobald. 
Hcemagogus^  Willis  ton. 


Ficalbia.  Theobald. 


UraiioicEnia^  Arribalzaga. 

Mi^nofnyia,  Theobald. 
^■Edeoinyia,  Theobald. 


Section  B. — Metanotum  ornamented  with  chastse, 

squamce  or  both. 
o.   With  chaetae  only. 

Proboscis  longer  than  whole  body  ;    lateral 

wing  scales  Tasniorhynchus-like    ... 
Proboscis  as  long  as  whole  body  in  2  ;  frons 

drawn  out  into  a  prominence  ;  wing  scales 

rather  broad  and  long 
Proboscis  not  as  long  as  the  whole  body  ; 

lateral  vein  scales  narrow  ... 
Proboscis  not  as  long  as  whole  body,  swollen 

apically  ;  wing  scales  long  and  broad 
)8.    Metanotum  with  squam.e  and  chsetce. 
Palpi  short  in  S^  and  ? . 
Proboscis  straight  in    $    and   S^  ;   legs  with 

scaly  paddles 

Venation  like  Sabethes. 

Legs  simple 

Venation  like  Culex 

Proboscis  in  ^  elbowed,  with  two  scaly  tufts 

Palpi  long  in  c^,  short  in  $ 

II.    Wings  with  seven-scaled  longitudinal  veins: 
Culex  type 


Phoniomyia^  Theobald. 

I  Binolia,  Blanch ard  = 
(  Runchiomyia,  Theobald. 

IVyeomyi'a,  Theobald. 
Dendriomyia^  Theobald. 

■Sabethes^  Rob.  Desvoidy. 

Sabethoides,  Theobald. 
Goeldia,  Theobald. 
Limattcs^  Theobald. 
Joblotina,  Blanchard. 

Heptaphlebo7nyia,  Theobald. 


Section    C. — Proboscis    short,    not    formed    for 
piercing 
Metatarsus  longer  than  first  tarsal  joint 
Metatarsus  shorter  than  first  tarsal  joint     ... 


Coreth^-ina. 
Corethra,  Linnaeus. 
Mochlonyx^  Ruthe.^ 


'  Many  other  genera  have  been  created  ;  these  will  be  fT)und  in  my  catalogue  of  Cidicidce 
ly   "Monograph  of  the  Mosquitoes  of   the  World,"   1901-10,    5  vols.,  in    my  "  Novse 


in  my 

Culicidae,"  family  CulicidiV 


Genera  Insectorum,  etc. 


66  THE   ANIMAL   PARASITES   OF   MAN 

Notes  on  the  Different  Genera. 
Sub-family.     Anophelina. 

The    following    Anophelines    have     been    recorded    as    malaria 
carriers  : — 

^^- Anopheles  maculipennis^  Meigen. 

Anopheles  bifurcatus,lAViv\dS:MS. 
* Myzomyia  funesta^  Giles. 

Myzomyia  lutzii,  Theobald. 
*Myzoinyia  rossii,  Giles. 

Myzomyia  listoftii^  Liston. 

Myzomyia  culicifacies^  Giles. 

Pyretophorus  superpictus,  Grassi. 
* Pyretophorus  costalis,  Loew. 

Pyretophorus  chaitdoyei,  Theobald. 
*Cellia  argyrotarsis^  Robineau  Desvoidy. 

Myzorhynchtts pseudopicitis^  Grassi. 

Myzorhynchus  barbirostris^  Van  der  Wulp. 

Myzorhynchus  sinensis^  Wiedemann. 

Myzorhynchus  paludis,  Theobald. 

Myzorhynchus  maiiritianus^  Grand  pre. 

Neocellia  stephensii,  Liston. 

Neocellia  willmori^  James. 

Nyssorhynchus  theobaldii^  Giles. 

Nyssorhynchus  fuliginosus^  Giles. 

Nyssorhynchus  annulipes,  Walker. 

Those  marked  with  an  asterisk  (*)  also  carry  the  larvae  of  Filaria  bancrofti^  as 
also  do  Myzorhynchus  minutus,  Theobald,  and  Myzorhynchus  Jiigerrinius,  Giles_ 

Genus.     Anopheles,  Meigen. 

"  Syst.  Beschr.  Europ.  zwei.  Ins.  I,"  1818,  ii,  p.  2,  Meigen  ;  "Mono.  Culicid.," 
1903,  i,  p.  191  ;  iii,  p.  17  ;  and  1910,  v,  p.  3,  Theobald. 

This  genus  contains  a  few  large  species  found  either  in  temperate  climates  or 
in  hills  and  mountains  of  warm  climates.  The  type  is  the  European  and  North 
American  A.  maculipennis. 


Fig.   399.  —  ^'xugoi  Atiophele'j  maciilipejinis,  MQ\^t\\. 

A.  7naculipennis,  Meigen.  This  species  and  A.  bifurcatus  are  malaria  carriers. 
True  Anopheles  only  occur  in  Europe,  North  America,  the  North  of  Africa  and  in 
the  mountains  of  India,  and  one  has  been  found  by  Bancroft  similar  to  A.  bifur- 
catus in  Queensland.  They  are  easily  told  by  the  absence  of  scales  on  thorax 
and  abdomen,  and  by  the  ratlier  densely  scaled  wings  with  lanceolate  scales. 


NOTES   ON   THE   DIFFERENT   GENERA  567 

Genus.     Myzomyia,  Blanchard  ;  Grassia,  Theobald. 

Coinp.  re?id.  heb.  Soc.  Biol.,  No.  23,  p.  795,  Blanchard  ;  "Mono.  Culicid.," 
1910,  iii,  p.  24  ;  V,  p.  16,  Theobald. 

This  genus  occurs  in  Asia,  Africa  and  South  America,  Europe  and  East  Indies. 
The  type  is  M.funesta^  Giles,  found  in  Central  and  West  Africa.  Although  structur- 
ally there  is  not  much  difference  between  this  genus  and  Anopheles,  they  differ 
greatly  in  appearance,  and  there  are  usually  a  few  narrow  curved  thoracic  scales 
projecting  over  the  head,  whilst  the  wing  scales  are  much  smaller  in  proportion,  and 
the  wings  more  uniformly  spotted,  always  so  along  the  costa.  Ftmesta  and  lutzii 
are  undoubtedly  malaria  bearers  and  also  rossii. 

Genus.     Neomyzomyia,  Theobald. 

"Mono.  Culicid., "1910,  v,  p.  29. 
A  single  species  only  occurs  in  this  genus,  N.  elegans^  James,  from  India.     In 
this  genus,  which  is  near  to  Myzomyia,  the  fourth  long  vein  is  very  near  the  base 
of  the  third,  and  there  are  outstanding  scales  on  the  prothoracic  lobes,  and  there  is 
a  marked  tuft  of  dense  scales  at  the  posterior  angles  of  the  head. 

Genus.     Cycloleppteron,  Theobald. 
"  Mono.  Culicid.,"  1903,  ii,  p.  312  ;  1903,  iii,  p.  58  ;  1910,  v,  p.  33. 

Two  common  species  only  occur  in  this  genus,  C.  grabhamii,  Theob.,  from 
Jamaica,  and  C.  mediopiinciatus^  Theob.  (Lutz.,  ms.),  from  South  America.  The 
chief  character  is  the  presence  of  large  black  inflated  pyriform  scales  on  the  wings. 
The  palpi  are  densely  scaled.     Neither  have  been  shown  to  be  malaria  bearers. 

Genus.     Feltinella,  Theobald. 
"  Mono.  Culicid.,"  1907,  iv,  p.  56. 
A  single  species,  so  far  only  found  in  this  genus.     The  basal  lobes  of  the  male 
genitalia  of  two  segments,  the  prothoracic  lobes  with  dense  outstanding  scales. 
The  species,  F.  pallidopalpi^  Theob.,  occurs  in  Sierra  Leone. 

Genus.     Stethomyia,  Theobald. 
"  Mono.  Culicid.,"  1903,  iii,  p.  13  ;  1907,  iv,  p.  59  ;  iQio,  v,  p.  35. 

Four  species  occur  in  this  marked  genus — one  S.  Jiimba,  Theob.,  from  British 
Guiana  and  Para,  another  S.  fiagalis,  Theob.,  from  the  Malay  States,  S.  culici- 
fonnis,  James  and  Liston,  from  India,  and  S.  pallida,  Ludlow,  from  India. 

The  former  may  be  a  malaria  carrier,  for  Dr.  Low  says  :  "  Malarial  fever  is  got 
amongst  the  Indians  and  often  of  a  severe  type.  In  that  connection  it  is  interesting 
that  in  the  interior,  at  a  place  called  Corato,  I  got  an  entirely  new  Anopheles  in 
large  numbers."  The  genus  is  easily  told  by  its  unornam-ented  wings,  flat  head 
scales,  mammillated  prothoracic  lobes  and  long  thin  legs. 

Genus.     Pyretophorus,  Blanchard;  Howardia,  Theobald. 

Compt.  rend.  heb.  Soc.  Biol.,  No.  23,  p.  705,  Blanchard  ;  Journ.  Trop.  Med.,v,  p.  181  ; 
and  "  Mono.  Culicid.,"  1903,  iii,  P-  13  ;  1910,  v,  p.  36,  Theobald. 
Forty-four  species  come  in  this  genus,  of  which  Anopheles  costalis,  Loew,  is  the 
type. 


^68  IHE   ANIMAL   PARASITES   OF   MAN 

This  genus  is  found  in  Africa,  India,  Europe  and  in  Australia.  Three  species 
are  proved  malaria  bearers,  namely,  P.  costalis,  Loew,  P.  chaudoyei,  Theob.,  and 
P.  superpictus,  Grassi.  Members  of  this  genus  can  be  told  by  having  narrow 
curved  thoracic  scales,  hairy  abdomen,  and  much-spotted  wings. 

Genus.     Myzorhynchella,  Theobald. 
"Mono.  Culicid.,"  1907,  iv,  p.  78. 
In  this  genus  the  thorax  has  distinct,  narrow  curved  scales,  and  the  abdomen 
is  hairy,  the  wing  scales  broad  and  lanceolate,  and  the  head  with  broad  scales  not 
closely  appressed,  but  not  forked  or  fimbriated. 

Five  species  are  known  :  lutziy  Cruz  ;  parva,  Chagas  ;  nigritarsis,  Chagas ; 
tibiomaculata,  Neiva ;  gilesi,  Neiva ;  and  nigra,  Theobald.  They  are  all 
recorded  from  Brazil,  and  7iigra  also  from   Mexico. 

Genus.     Manguinhosia,  Cruz,  in  Peryassu. 

•*Os  Culicideos  do  Brazil,"  1908,  p.  112. 

A  single  marked   species    from    the  Brazils.     The  thorax  has  piliform  curved 

scales,  and  some  narrow  curved  and  flattened  ones  on  the  sides.     Abdomen  pilose, 

except  the  last  three  segments  which  are  scaled.     No  tufts  of  scales  on  posterior 

femora. 

Allied  to  Lophoscelomyia,  but  at  once  told  by  the  absence  of  scale  tufts  on  the 
hind  femora.     M,  lutzi,  Cruz,  Brazil. 

Genus.     Chrystya,  Theobald. 

"  Rep.  Sleeping  Sickness,  Roy.  Soc.  Eng.,"  1903,  vii,  p.  34. 
A  very  marked  genus  in  which  the  hairy  abdomen  has  very  long,  dense,  hair-like, 
apical,  scaly  tufls  to  the  segments.     A  single  species  only  so  far  known,  C.  implexa^ 
Theobald,  from  Africa  (Uganda,  Sudan,  etc.). 

Genus.     Lophoscelomyia,  Theobald. 

Entomologist^  1904,  xxxvi,  p.  12. 
A  single  species  only,  from  the  Federated  Malay  States.    The  hind  femora  have 
dense,  apical  scale  tufts  ;  the  thorax  long,  hair-like  curved  scales  ;  abdomen  pilose, 
except  the  last  two  segments  which  are  scaly  ;  wings  with  broad,  blunt,  lanceolate 
scales. 

Genus.     Arribalzagia,  Theobald. 

"  Mono.  Culicid.,"  1903,  iii,  pp.  13  and  81  ;  and  1910,  v,  p.  48. 
Two  species  only  occur,  found  in  South  America.  The  thorax  and  abdomen 
have  scales  and  hairs  respectively,  as  in  Fyretophorus,  but  the  abdomen  has  in 
addition  prominent  lateral  apical  scale  tufts  to  the  segments  and  a  scaly  venter. 
Wings  with  membrane  tinged  in  patches  and  wing  scales  bluntly  lanceolate  and 
very  dense.  The  type  is  A.  maculipes,  Theob.  found  in  Trinidad  and  Brazil; 
A. pseudoinaculipes^  Cruz,  also  in  Brazil, 

Genus.     Myzorhynchus,  Blanchard  ;   Rossia,  Theobald. 
Compt.  rend.  heb.  Soc.   Biol.,   1902,  No.   23,  p.   795,  Blanchard  ;  Journ.   Trop. 
Med.,  1902,  p.  181,  Theobald  ;  "Mono.  Culicid.,"  1903,  iii,  p.  84;   1907,  iv,  p.  80; 
1 9 10,  V,  p.  49. 

A  very  marked  genus  of  large,  dark,  densely  scaled  species,  found  in  Europe, 
Asia,  Africa  and  Australia.     The  thorax  with  hair-like  curved  scales;  the  abdomen 


NOTES   ON   THE   DIFFERENT   GENERA  5O9 

with  ventral  and  apical  scales,  and  a  median  ventral  apical  tuft,  and  with  very 
densely  scaled  palpi  in  the  female,  and  densely  scaled  proboscis.  It  seems  to  be 
mainly  an  Asiatic  and  East  Indian  genus,  but  three  species  occur  in  Africa  and 
one  in  Australia.     They  are  mostly  sylvan  species  and  bite  severely. 

Fourteen  species  are  known.     Five  are  malaria  carriers  {vide  list,  p.  566). 

Genus.     Nyssorhynchus,  Blanchard  ;  Laverania,  Theobald. 

"  Mono.  Culicid.,"  1910,  iii,  p.  14  ;  v,  p.  55,  Theobald  ;  Compt.  rend.  heb.  Soc.  Biol.^ 
No.  23,  p.  795,  Blanchard. 

A  group  of  small,  closely  allied  species  found  in  Asia,  Africa  and  Australia, 
twelve  out  of  the  twenty  species  coming  from  India. 

The  thorax  is  covered  with  narrow  curved  and  spindle-shaped  scales,  abdomen 
with  small,  flat  or  narrow  curved  dorsal  scales,  especially  on  the  apical  segments  or 
in  patches  ;  the  legs  are  always  banded  or  spotted  with  white,  and  the  tarsi  have  as 
a  rule  one  or  more  pure  white  segments.  •  (This  banding  and  spotting  is  of  no 
generic  value,  however.) 

The  species  show  considerable  seasonal  variation.  The  type  of  the  genus  is 
N.  maculaius^  Theobald. 

Three  are  malaria  carriers  {vide  list,  p.  566). 

Genus.      Cellla,  Theobald. 

"  Mono.  Culicid.,"  1903,  iii,  p.  107  j  1910,  v,  p.  67. 

Very  marked  Anophelines,  with  densely  scaly  abdomens,  the  scales  irregularly 
disposed  on  the  dorsum  and  forming  dense  lateral  tufts  ;  thorax  with  flat  spindle- 
shaped  scales  ;  palpi  densely  scaled  and  also  the  wings. 

The  type  of  the  genus  is  the  African  C. pharoensis^  Theob.  It  is  represented  in 
Asia  by  C.  kochii,  Donitz  ;  in  West  Indies  and  South  America  by  C.  argyrotarsis , 
Desvoidy,  and  C.  bigotii,  Theob. ;  in  Africa  by  C.  squamosa^  Theob.,  etc. 

C.  argyrotarsis ,  Desvoidy,  and  C.  albimana,  Wiedemann,  are  undoubtedly 
malaria  bearers. 

Genus.     Neocellia,   Theobald. 

"Mono.  Culicid.,"  1907,  iv,  p.  in. 
Allied  to  Cellia,  but  has  no  lateral  scale  tufts.     Three  species  recorded  froui 
India. 

Genus.      Kert^szia,  Theobald. 
"Ann.  Mus.  Nat.,  Hung.,"  1905,  iii,  p.  66. 
This  genus  has  the  thoracic  scales  hair-like,  except  a  few  narrow  curved  ones 
in  front ;  abdominal  scales  long,  broad  and  irregular. 

A  single  species,  K.  boliviensis,  Theob.  from  Bolivia. 

Genus.      Manguinhosia,  Cruz. 

The  thorax  has  narrow  hair-like  curved  scales  and  some  broad  straight  scales  ; 
others  spatulate  on  the  sides.  Abdomen  with  fine  hairs,  except  the  last  three 
segments  which  are  scaly.  Tufts  of  scales  on  the  hind  femora.  Wing  scales 
lanceolate. 

The  type  is  M.  lutzi,  Cruz,  from  Brazil. 


cyo  THE   ANIMAL   PARASITES   OF   MAN 

Genus.     Chagasia,  Cruz. 

"Brazil-Medico,"  1906,  xx,  pp.  20,  199. 
This  genus  can  at  once  be  told  by  the  antennal  segments  having  many  dense 
scaly  tufts.     Type,  C.fajardoi,  Lulz,  from  Brazil. 

Genus.     Calvertina,  Ludlow. 

Canadian  Entomologist,  1909,  xli,  pp.  22,  234. 
The  antennae  in  this  genus  have  outstanding  scales  on  the  second  segment,  more 
appressed  ones  on  the  first.     At  least  one  abdominal  segment  with  long,  flat,  more 
or  less  spatulate  scales.     Type,  C.  lineata,  Ludlow,  from  Philippine  Islands. 

Genus.     Birdnella,  Theobald. 

"Ann.  Mus.  Nat.  Hung.,"  1905,  iii,  p.  69. 
At  once  told  by  the  first  submarginal  cell  being  very  small.     Type,  B.  gracilisy 
Theob.  from  New  Guinea. 

Sub-family.     Megarhininae. 

Three  genera  occur  in  this  marked  sub-family ;  they  are  the  largest  of  all  mos- 
quitoes, and  are  very  brilliantly  coloured,  and  many  have  tail  fans.  They  occur 
in  North  and  South  America,  Asia,  Africa,  and  Australia.  The  long  curved 
proboscis  is  very  marked.  They  are  usually  spoken  of  as  elephant  mosquitoes  ; 
some  are  vicious  blood-suckers  at  times. 

The  three  genera  tabulate  as  follows  : — 
a.    Palpi  long  in  both  sexes. 
&.   Last  segment  of  $  palp  round  or  blunt 

as  if  broken       Genus  A/if^^^r/zZ/mi-,  R.  Desvoidy. 

)8;8.  Last   segment  of   $    palp   long   and 

pointed         Genus  Aiikylorhyjichus,  Lutz. 

00.  Palpi  of  female  short  of  male  long. 
Palpi  of  female  not  more  than 
one-third  length  of  proboscis       ...     Q^xvws  roxorhyfichites,T\i^o\i2\d.. 

Genus.     Megarhinus,  Robineau  Desvoidy. 

"  Mem.  Soc.  d'Hist.  nat.  de  Paris,"  1827,  iii,  p.  412  :  "  Mono.  Culicid.," 
1901,  i,  p.  215  ;  1903,  iv,  p.  163;  1907,  iv,  p.  128  ;  1910,  v,  p.  89. 
All  large  brilliant  mosquitoes  with  long  palpi  in  both  sexes  and,  as  a  rule,  with 
a  caudal  fan  of  scales  ;  the  proboscis  is  long  and  bent.     They  are  all  sylvan  species, 
and  are  not  so  far  recorded  as  biting  man. 

Genus.     Toxorhynchites,  Theobald. 
"  Mono.  Culicid.,"  1901,  i,  p.  244  ;  1903,  iii,  p.  1 19  ;  1907,  iv,  p.  140 ;  1910,  v,  p.  95. 
Differs  from  the  former  genus  in  that  the  female  palpi  are  short.    The  palpi  may 
have  one,  two  or  three  minute  terminal  segments.     Banks's  genus  Worcesteria  has 
three. 

The  elephant  mosquito  of  India  {T.  iimnisericors),  Walker,  bites  very  severely. 
They  are  sylvan  species. 


NOTES   ON   THE    DIFFERENT   GENERA  571 

Sub-family.     Culicinae. 
Genus.    Mucidus,  Theobald. 

"Mono.  Ciilicid.,'"  1901,  i,  p.  268  ;  1910,  v,  p.  125. 

This  genus  is  so  far  confined  to  Australia,  West  and  Central  Africa,  India,  East 
Indies  and  Malay  Peninsula.  They  are  all  large  mosquitoes,  easily  told  by  the 
whole  body  being  more  or  less  covered  with  long  twisted  scales,  giving  them  a 
mouldy  appearance,  and  the  legs  densely  scaled  with  outstanding  scales  ;  the  wings 
.with  large  parti-coloured  scales.  The  Australian  M.  a//ernans,  Walker,  occurs  in 
larval  form  both  in  fresh  and  salt  water.     The  adults  bite  man. 

Genus.     Psorophora,  Robineau  Desvoidy. 

"  Mem.  de  la  Soc.  d'Hist.  nat.  de  Paris,"  1827,  iii,  p.  412,  R.  Desvoidy  ;  "  Mono. 
Culicid.,"  1901,  i,  p.  259;  1903,  iii,  p.  130;  1907,  iv,  p.  158  ;  i9io,v,  p.  123,  Theobald. 

This  genus  is  confined  to  the  Americas  and  the  West  Indies.  Several  species 
exist  which  can  easily  be  told  from  Mucidus  by  the  absence  of  long  twisted  scales 
and  the  narrower  wing  scales.  The  legs  are  densely  scaled  and  the  thorax  orna- 
mented with  flat  spindle-shaped  scales. 

F.  ciliata^  Robineau  Desvoidy,  occurs  in  both  North  and  South  Americfj,  and 
bites  man. 

Genus.     Janthinosoma,  Arribalzaga. 

"Dipt.  Arg.,'  1891,  p.  52,  Arribalzaga;  "Mono.  Culicid.,"  1901,  i,  p.  253  ; 
1903,  iii,  p.  124  ;   1907,  iv,  p.  152  ;  and  1910,  v,  p.  118,  Theobald. 

Hind  legs  only  densely  scaled  ;  some  of  the  hind  tarsi  are  always  white.  The 
venation  is  as  in  Culex.  The  abdomen  is  metallic  and  iridescent.  They  all  bite 
man  and  occur  only  in  the  Americas  and  West  Indies. 

Genus.      Stegomyia,  Theobald. 

"  Mono.  Culicid.,"  1901,  i,  p.  283  ;  1903,  iii,  p.  130  ;  1907,  iv,  p.  170;  1910,  v,  p.  151. 

This,  the  most  important  genus  in  the  Culicince^  can  be  told  by  the  head  and 
scutellum  being  clothed  with  flat  scales  and  the  thorax  with  narrow  curved  ones. 

About  forty  species  are  known  in  this  genus,  occurring  in  Southern  Europe,  Asia, 
Africa,  Australia,  the  Americas,  East  and  W^est  Indies,  and  on  most  oceanic  islands. 
Many  of  them  seem  to  be  vicious  blood-suckers.     They  are  mostly  black  and  white 
mosquitoes,  and  several  seem  to  go  by  the  name  of  tiger  mosquitoes.     The  genus 
contains  the  yellow  fever  mosquito  (6*.  fasciata^  Fabricius),  the  only  one  that  need 
be  dealt  with  in  detail  here.     The  chief  known  species  tabulate  as  follows  : — 
A.  Proboscis  banded. 

a.   Legs  basally  banded. 

Thorax  brown,  with  scattered  creamy-white 

scales  ...         ...         ...         ...         ...     annulirostris^  Theobald. 

Thorax  black,  with  narrow,  curved  golden 

scales  ...         ...         ...         ...         ...    periskelta^  Giles. 

«o.  Legs  with  basal  and  apical  banding.  Fore 
legs  with  no  bands  ;  mid  with  apical  and 
basal  bands  on  first  and  second  tarsals, 
hind  with  basal  bands. 
Thorax  white  in  front,  with  a  brown  eye- 
like spot  on  each  side        ///i?;;/^^?///,  Theobald. 

36 


572 


THE   ANIMAL   PARASITES   OF   MAN 


AA.  Proboscis  unhanded. 
/3.  Legs  basally  banded. 
%  Abdomen  basally  banded. 

Thorax  with  one  median  silvery-white  line 

Thorax  as  above,  but  pleurae  with  white 
lines 

Thorax  similar,  but  two  white  spots  near 
where  line  ends       

Thorax  with  two  median  yellow  lines  and 
lateral  curved  silvery  lines 

Thorax  with  two  short  median  lines  and  a 
white  patch  on  each  side 

Thorax  with  large  lateral  white  spots  in 
front,  smaller  ones  by  wings,  two  narrow 
median  lines  and  two  posterior  sub- 
median  white  lines... 

Thorax  with  a  white  W-shaped  area  in 
front,  a  prolongation  curved  on  each  side 
enclosing  a  brown  eye-like  spot  ... 

Thorax  with  white  frontal  median  spot,  two 
large  lateral  spots,  a  small  one  in  front  of 
the  wings,  a  narrow  median  white  line 
and  narrow  sub-median  ones  on  posterior 
half.     Last  two  hz?id  tarsi  white  ... 

Thorax  brown,  with  broad  white  line  in 
front  extending  laterally  towards  wings, 
where  they  swell  into  a  large  patch,  a 
white  line  on  each  side  just  past  wing 
roots.     Last  two  hind  tarsi  white 

Thorax  with  silvery  white  spot  on  each  side 
in  front,  small  one  over  roots  of  wings 
and  white  over  their  base.  Last  two  hind 
tarsi  white 

Thorax  with  two  lateral  white  spots,  front 
ones  the  largest,  a  small  median  one  near 
head,  two  yellow  median  lines,  a  short 
silvery  one  on  each  side  before  the 
scutellum      

Thorax  with  silvery  white  scaled  area  in 
front  and  another  on  each  side  in  front  of 
wings  

Thorax  with  median  yellowish-white  line,  a 
silvery  patch  on  each  side  in  front  of 
wings  extending  as  a  fine  yellow  line  to 
the  scutellum,  and  another  silvery  spot 
before  base  of  each  wing 

Thorax  with  small  grey-scaled  area  in  front 
of  wing  roots  and  three  short  creamy  lines 
behind 


scutellaris^  Walker. 
Pseudosctitellaris^  Theobal d. 
geleln7tensis,  Theobald. 
fasciata^  Fabr. 
nigeria^  Theobald. 

Ulii^  Theobald. 
W-alba,  Theobald. 

wellinannii^  Theobald. 

albipes^  Theobald. 
pseudonigeria,  Theobald. 

simpsoni,  Theobald. 
argenteojnaculata^  Theobald. 

poweri^  Theobald. 
ininutissima^  Theobald. 


NOTES   OX   THE    DIFFERENT   GENERA 


573 


Thorax  (?)  denuded  ;  abdomen  black  ;  fifth 
segment  with  yellow  basal  band  ;  sixth 
unhanded  ;  seventh,  two  median  lateral 
white  spots  ;  eighth,  two  basal  lateral 
white  spots  ;  second  hind  tarsal  nearly  all 

white  ...         

77.  Abdomen  unhanded. 

First  hind  tarsal  all  white,  second  basally 
white,  last  two  dark.  Thorax  chestnut 
brown,  with  a  broad  patch  of  white  scales 
on  each  side  in  front  and  a  median  pale 

line 

80.  Legs  with  white  lines  as  well  as  basal  bands. 
Thorax  brown,  with  white  lines  ;  abdomen 

with  basal  bands 

000.  Fore  and'mid  legs  with  apical  bands,  hind 

basal. 

Fourth  tarsal  of  hind  legs  nearly  all  white 

Mid  metatarsi  with  basal   pale  banding, 

base  and  apex  of  hind,  also  base  of  first 

tarsal  pale 

0000.  Legs  unhanded. 

5.  Abdomen  basally  banded. 

Thorax  with  front  half  white,  rest  bronzy- 
brown  ...         ...         

Thorax      deep      brown,     with     scattered 
golden    scales,  showing  two   dark    eye- 
like   spots  ;    head    white,  dark  on  each 
side  and  behind  ... 
Thorax     brown      with      golden 
abdomen    with    narrow  basal 
fifth  and  sixth  segments  only     ... 
55.  Abdominal  banding  indistinct. 

Thorax  with    broad    silvery    white 
on  each  side  in  front 
?55.  Abdomen  unhanded. 

Thorax  with  six  silvery  spots 

Abdomen  with  apical  white  lateral  spots. 

Thorax  unadorned,  except  for  pale  scaled 

lines  laterally 
Abdomen  with  basal  white  lateral  spots. 
Thorax    with   two    pale    median    parallel 

lines  and  two  silvery  lateral  spots 
Thorax  unadorned. 

A  white  spot  middle  of  head  ... 
No  white  spot  ... 
AAA.  Proboscis  yellow  basally,  dark  apically. 
Abdomen  with  apical  pale  bands  ... 
A  \AA.  Proboscis  with  median  interrupted   white 
line  on  basal  half. 
Head  black,  anterior  margin  grey 


2555. 


stripes  ; 
bands  on 


patch 


55355. 


{/uh'a,  Theobald. 


terreus.,  Walker. 
grantii^  Theobald. 
mediopunctata.,  Theobald. 
assamemis,  Theobald. 

pseudonivea,  Ludlow. 

albocep/iala^  Theobald. 
nuriostrjaia.,  Banks. 

albolateralis.,  Theobald. 
argenteopmtcfafa,  Theobald. 

punctolateralis.,  Theobald. 

iiiinuta,  Theobald. 

tripunctata.,  Theobald. 
amesii,  Ludlow. 

crassipes,  Van  der  Wulp. 
albomarginata^  Newstead. 


574 


THE   ANIMAL   PARASITES    OF    MAX 

Stegomyia  fasciata,  Fabriciiis  (Yellow  Fever  Mosquito). 


This  insect,  which  is  the  proven  carrier  of  yellow  fever,  is  commonly  called  the 
tiger,  brindled,  spotted  day  or  striped  mosquito.  It  is  also  referred  to  by  some 
writers  as  S.  ca/opus,  Meigen.  It  is  subject  to  considerable  variation  in  colour, 
but  the  thoracic  markings  are  generally  very  constant.  The  general  colour  is 
almost  black  to  deep  brown,  the  head  with  a  median  white  area,  white  at  the  sides 
and  in  front  around  the  eyes  ;  the  thorax  has  two  median  parallel  yellow  lines, 
a  broad  curved  silvery  one  on  each  side  and  white  spots  at  the  sides  ;  tlie  scales  on 
the  intervening  spaces  of  the  thorax  are  brown.  The  dark  abdomen  has  basal  white 
bands  and  basal  white  lateral  spots.  The  dark  legs  have  basal  white  bands,  the 
last  segment  of  the  hind  legs  being  all  white  except  in  a  variety  from  South  America 
and  the  West  Indies  {luciensis),  which  has  the  tip  of  the  last  hind  tarsal  dark.  The 
abdomen  may  also  vary  in  colour,  some  having  pale  scales  over  most  of  the  surface 
{queenslandetisis). 

The  food  of  the  adult  female  consists  mainly  of  man's  blood,  but  she  will  also 
feed  on  dogs  and  other  animals.  The  male  has  been  said  to  bite,  but  such  is  very 
unusual.     This  mosquito  bites  mainly  in  the  daytime  up  till  about  5  p.m. 

The  adults  breed  the  first  day  after  emergence.  They  may  live  a  consider- 
able time,  Bancroft  having  kept  females  for  two  months  in  confinement.  The 
ova  are  laid  separately,  often  in  chains ;  they  are  black,  oval,  with  a  reticulated 
membrane  outside,  some  of  the  reticulated  cells  containing  air.  They  may  hatch 
in  from  six  to  twenty  hours,  the  larval  stage  nine  days,  the  pupal  stage  three  ;  thus 
the  whole  cycle  may  be  completed  in  from  twelve  to  thirteen  days.  The  ova  when 
dry  can  remain  undeveloped  for  a  considerable  time.  The  larvae  are  greyish-white, 
with  short,  thick  siphon,  and  feed  at  the  bottom  of  the  water,  only  coming  to 
the  surface  now  and  again  to  breathe.  This  is  almost  entirely  a  domesticated 
gnat,  seldom  being  found  far  from  man's  habitations.  Its  larvne  occur  in  such 
small  collections  of  water  as  old  sardine  tins,  jam-pots,  calabashes,  puddles,  barrels, 
wells — in  fact,  wherever  water  is  held  up,  even  to  the  gutters  of  houses.  Not 
only  are  they  found  breeding  on  land,  but  also  on  board  ship,  although  they 
prefer  artificial  collections  of  water.  They  may  also  breed  in  larger  natural 
collections. 

This  insect  is  easily  transported  by  steam  and  sailing  ships  and  by  train,  and 
this  doubtless  explains  its  very  wide  distribution.  The  adults  may  live  for  fifty 
days,  and  it  is  on  this  account  and  their  frequent  occurrence  on  ships  that  danger 
lies  in  regard  to  the  Panama  Canal.  An  infected  insect  may  leave  that  endemic 
centre  of  yellow  fever  and  live  until  the  vessel  arrived  at  the  Philippine  Islands  and 
fly  ashore,  and  so  introduce  the  disease  for  the  wdi^w^  fasciata  possibly  to  spread. 

Roughly  the  distribution  of  this  pest  is  as  follows  :  Africa  from  South  to  North, 
but  especially  along  the  coast  and  up  the  Nile.  In  Asia,  in  India,  Ceylon,  Burma, 
Siam,  along  the  ports  of  the  Malay  Peninsula,  in  French  Cochin  China,  Philippine 
Islands,  the  Andaman  and  Nicobar  Islands,  Japan,  Malay  Archipelago,  and  East 
Indies,  Turkey  in  Asia,  Arabia  and  Palestine. 

In  Australia  it  occurs  in  Queensland,  New  South  Wales,  Victoria  and  South 
Australia. 

In  Europe  in  Italy,  Spain,  Portugal,  Greece,  in  the  Mediterranean  Islands. 

In  South  America,  Central  America,  Mexico,  North  America,  and  the  W^est 
Indies  it  is  very  abundant,  and  it  also  is  found  in  the  Bahama  Islands,  Fiji,  Sandwich 
Islands,  Samoa,  the  Azores,  Teneriffe  and  Santa  Cruz,  Pitcairn  Islands  and  Bermuda. 
For  a  full  account  of  its  distribution  the  reader  is  referred  to  the  following  : 
"The  Distribution  of  the  Yellow  Fever  Mosquito  {Stegomyia  fasciata^  Fabricius) 
and    General    Notes     on    its    Bionomics;"     "Mem.    le""    Congres    international 


THEOBALDIA,    CULEX  575 

d'Entomologie,  191 1,  ii,  pp.  145-170,  F.  V.  Theobald.  In  addition  to  being  the 
yellow  fever  carrier,  it  is  supposed  by  Wenyon  to  be  the  intermediate  host  of  the 
parasite  of  Bagdad  sore. 

Stegomyia  scutellaris,  Walker. 

A  vicious  biter,  found  in  India,  China,  Malay,  East  Indies,  and  Ceylon.  The 
thorax  has  one  median  silvery  stripe,  and  so  can  easily  be  told  from  S.fasciata. 

A  very  similar  species  occurs  in  Fiji,  but  can  be  told  by  the  pleurae  having  white 
lines,  not  spots  {S.  pseudoscutellaris^  Theobald).  It  is  the  intermediate  host  of 
filaria  in  Fiji  (Bahr). 

A  number  of  nearly  allied  genera  occur  here  (c^'ide  synoptic  table). 

Genus.     Theobaldia,   Neveu-Lemaire. 
Theobaldinella,    Blanchard. 

Includes  several  large  Culicines,  of  which  T.  a?tmilata^  Meigen,  is  the  type.  The 
wings  are  usually  spotted  {aniiulata^  i?tcidens,  etc.),  but  may  be  nearly  plain  {spathz- 
palpis).  The  males  have  the  palpi  swollen  apically,  and  the  females  have  long 
five-jointed  palps. 

Several  of  these  are  vicious  biters. 

Theobaldia  annulata,  Meigen. 

This  large  gnat  (6  mm.  long)  can  be  told  by  its  wings  having  five  large  spots  of 
dark  scales  and  by  its  legs  having  broad  basal  white  bands  to  the  tarsi.  The  larvas 
occur  in  rain  barrels  and  small  pools.  It  is  essentially  a  domestic  form,  occurring  in 
houses  and  privies.  Its  distribution  is  Europe  generally  and  North  America.  The 
bite  is  very  severe,  and  in  some  districts  gives  rise  to  painful  oedema.' 

Theobaldia  spathipalpis,  Rondani,  occurs  in  Italy,  Mediterranean  Islands, 
Palestine,  the  Himalayas,  Khartoum,  and  in  South  Africa.  It  is  about  the  same  size 
as  T.  annulata,  but  is  yellowish-brown  in  colour,  with  striped  thorax  and  mottled  and 
banded  legs.     It  occurs  in  privies  and  bites  very  severely. 

Genus.     Culex,  Linnaeus. 

"  Syst.  Nat.  Ed.,"  1758,  x,  Lmn.neus  ;    "Mono.  Culicid.,"  1901,  i,  p.  326  ; 
1910,  V,  p.  322,  Theobald. 

This  large  genus  still  contains  many  forms  which  should  be  excluded.  The 
species  normally  have  narrow  curved  median  head- scales,  and  similar  ones  on  the 


Fig.  400.— Wing  of  a  Culex. 

scutellum  ;   the   female   palpi  are  shorter  than  in  the  former  genus  and  the  male 
palpi  are  pointed  ;  the  lateral  vein-scales  are  narrow  and  linear. 

The  type  is  Culex  pipie?ts,  Linn.,  the  common  gnat  of  Europe.     The  thorax 


Theobald,  "Second  Report  on  Economic  Zoology,"  1903,  p.  9. 


576  THE   ANIMAL   PARASITES   OF   MAN 

is  covered  with  narrow  curved  golden-brown  scales,  the  abdonien  has  basal  pale 
bands  to  the  segments  and  the  legs  and  proboscis  are  unhanded.  The  stem  of  the 
first  submarginal  cell  is  always  less  than  one-fifth  the  length  of  the  ceil.  It  lays  its 
eggs  in  rafts  in  water-butts,  etc.,  and  even  in  the  foulest  water.  They  are  first 
deposited  in  England  in  June  and  July,  and  again  soon  after  hatching  in  August.  In 
some  districts  this  gnat  bites  man  viciously,  in  others  not  at  all. 

The  common  tropical  gnat  {Culex  fatigans,  Wied).  This  resembles  the 
European  Culex  pipiens,  but  can  always  be  told  by  the  stem  of  the  first  submarginal 
cell  always  being  much  longer  than  it  is  in  C.  pipie?ts.  This  is  one  of  the  species 
that  has  been  proved  to  transmit  filariae  to  man,  etc.  Varieties  of  it  occur  in  almost 
every  country  between  40°  N.  and  S.,  having  a  very  similar  range  to  S.  fasciata.  In 
all  countries  it  appears  to  be  connected  with  the  transmission  of  Filaria  bancrofti^ 
and  it  is  also  said  to  carry  the  micrococcus  of  dengue  fever. 


Genus.     Melanoconion,  Theobald. 

"  Mono.  Culicid.,"  1903,  iii,  p.  238  ;   1907,  iv,  p.  507  ;   1910,  v,  p.  455. 

This  genus  is  composed  of  eight  species,  most  of  which  are  small  black  gnats 
which  bite  viciously  and  which  occur  in  swamps  and  jungles.  They  can  at  once 
be  told  from  Culex  by  the  veins  of  the  wings  having  dense  broadened  scales  on  their 
apical  areas  and  along  the  upper  costal  border.  The  femora  and  apices  of  the  tibiae 
are  swollen. 

The  black  mosquito,  Melanoco7iion  atraius,  Theob.  This  small  gnat  is  a  very 
troublesome  pest  in  swamps  in  the  West  Indies.  The  female  bites  both  by  day  and 
by  night,  and  the  bite  causes  severe  irritation.  The  larvie  live  in  permanent  ponds. 
It  is  almost  black  in  colour,  but  sometimes  presents  a  dull  coppery  sheen  ;  each 
segment  has  small  lateral  basal  white  spots.     Length  2*5  to  3  mm. 

It  occurs  in  Para  and  British  Guiana  as  well  as  in  the  West  Indies. 

Ordinary  mosquito  netting  is  no  use  for  keeping  off  this  pest. 


Genus.     Grabhamia,  Theobald. 

"  Mono.  Culicid.,"  1903,  iii,  p.  243.;  1907,  iv,  p.  284  ;  and  1910,  v,  p.  277. 
Allied  to  Culex,  but  separated  by  the  wings  having  short  fork-cells,  mottled  scales, 
the  median  ones  thick  and  also  some  of  the  lateral  ones  short  and  broad  ;  the  last 
two  joints  of  the  male  palps  are  very  slightly  swollen.  The  eggs  are  laid  singly,  not 
in  rafts,  and  the  larvae  have  short,  thick  siphons.  Ten  species  occur  and  are  found 
in  Europe,  North  America,  West  Indies  and  Natal.  G.  dorsalts,  Meigen,  bites 
severely  in  Europe.  G.  sollicitans.  Walker,  is  a  great  scourge  along  the  New  Jersey 
Coast  and  at  Virginia  summer  resorts  and  in  Florida.  It  breeds  in  brackish  water 
and  is  the  most  common  mosquito  of  the  Atlantic  seaboard. 


Genus.     Pseudotaeniorhynchus,  Theobald  ; 
Taeniorhynchus,     Theobald,     non-Arribalzaga. 

Differs  from  the  former  in  having  the  whole  wing  veins  clothed  with  dense, 
broadish  elongated  scales.  1  hey  occur  in  South  America  {T.  fasciolatus,  Arri.),  in 
Africa  (T.  tenax,  Theob.),  in  Europe  {T.  richardii,  Ficalbi).  The  latter  bites  very 
severely. 


OTHER   NEMATOCERA  577 

Genus.     Taeniorhynchus,  Arribalzaga  ;  Mansonia,  Blanchard  ; 
Panoplites,  Theobald. 

Compt.  rend.  heb.  Soc.  Biol.,  1901,  iii,  -^j^  p.  1046;  "Mono.  Culicid.," 
1901,  ii,  p.   173;  and  1910,  v,  p.  446,  Theobald. 

A  very  marked  genus,  easily  told  by  the  broad  asymmetrical  wing  scales.  It 
occurs  in  Africa  {T.  africana  and  T.  major,  Theob.)  ;  in  Asia  {T.  uniformis,  Theob. ; 
T.  annuHpes,  Walker,  etc.)  and  in  Australia  (7".  aiistraliensis)  ;  in  the  Americas  and 
West  Indies  (7".  titillans,  Walker).  The  eggs  (fig.  395,  d)  are  peculiar  in  form  and 
are  laid  separately ;  the  larva  has  not  been  described  ;  the  pupa  has  long  curved 
siphons.  They  mostly  occur  along  rivers,  in  swamps  and  forests,  and  bite  very 
severely.  They  also  enter  houses  {T.  titillans).  T.  imiforinis  is  most  troublesome 
during  the  rains.  The  saliva  is  strongly  acid.  Both  these  species  carry  the  larvae  of 
Filaria  bancrofti. 

Genus.     Chrysoconops,  Goeldi. 

"  Os  Mosq.  no  Para,"  1905,  p.  114,     Goeldi;   "Mono.  Culicid.,"   1910, 
V,  P-  433,  Theobald. 

Bright  yellow  or  yellow  and  purple  mosquitoes,  with  rather  dense  wing  scales. 
Numerous  species  occur  in  Africa  {aurites,  amtettii,  fuscopentiatus,  etc.),  others  in 
India,  Australia  and  South  America. 

Low  found  filarias  in  the  thoracic  muscles  o^  fuscopennatus  in  Uganda. 

Several  of  the  Aideojuyina  bite,  especially  the  small  Uranotccnias.  They  are  all 
sylvan  species,  seldom  entering  houses.    They  need  not,  therefore,  be  referred  to  here. 

For  full  details  of  the  Culicid  genera  and  species  the  reader  is  referred  to  my 
monograph'  and  other  works  mentioned  below. 

Other  Nematocera. 

Other  nematocerous  flies  are  midges,  daddy-long-legs  and  sand-flies.  The  ones 
which  cause  annoyance  to  man  besides  Cidicidce  are  the  followmg  : — 

Sand-flies  {Simididcr),  certain  midges  {Chironoinidce),  and  a  few  owl  midges 
{Psychodidcp). 

The  Neinatocera  have  long  thread-like  jointed  antennas  and  their  pupae  are,  as  a 
rule,  naked  ;  the  larvae  have  a  distinct  head  and  can  thus  be  told  from  the  next 
section  {Brachycerd). 

Family.     Simulldae. 

This  family  consists  of  a  single  genus,  Simulium,  Latreille,  which  Roubaud  has 
recently  divided  into  two  sub-genera  called  Pro-Simulium  and  Eu-Simulium.  These 
insects,  which  are  frequently  spoken  of  as  sand-flies,  are  found  in  all  parts  of  the 
world;  they  are  all  small  insects  varying  from  1-5  to  3  mm.  The  females  are  very 
bloodthirsty,  but  the  males  appear  to  be  incapable  of  sucking  blood. 

The  head  sunk  under  the  humped  thorax  ;  antennae  short,  straight ;  palpi  short 

»  **  A  Monograph  of  the  Cnlicidtt  of  the  World,"  5  vols,  and  atlas,  1901  to  1910,  British 
Museum  (Nat.  Hist.)  ;  and  the  following  :  Howaid,  Dyar  and  Knab,  "The  Mosquitoes  of 
North  and  Central  America  and  the  West  Indies,"  1912 ;  James  and  Liston,  "The 
AnophelinK  of  India,"  Leicester,  1908;  "The  Culicidcc  of  Malay,"  Inst.  Med.  Res.,  Fed. 
Malay  States,  iii ;  Ann.  Trop.  Med.  and  Par.,  papers  by  Ne\\  stead  and  Carter;  Mem,  Inst. 
Oswaldo  Cruz,  papers  by  Luiz,  Neva,  Chagas  ;  and  the  Bulletin  of  Entomological  Research,  etc 


578  THE   ANIMAL   PARASITES   OF   MAN 

and  broad,  of  four  segments,  bent ;  wings  broad  and  in  some  iridescent,  legs  stout. 
The  male  has  holoptic  eyes,  whilst  in  the  female  they  are  small  and  widely  separate. 
The  sucking  proboscis  is  short.  The  thorax  and  abdomen  are  clothed  with  short 
hairs  which  may  form  spots  and  markings ;  these  are  golden,  silvery,  grey,  or  brown- 
ish. In  the  sub-genus  Pro-Simulium  the  second  segment  of  the  hind  tarsi  in  both 
sexes  is  elongate,  linear,  and  without  a  basal  notch  ;  in  Eu-Simulium  it  is  short, 
curved,  and  dorsally  notched  at  the  base. 

Simulidce  often  occur  in  swarms,  and  attack  not  only  man  but  cattle,  horses, 
and  poultry.     In  some  districts  they  are  more  annoying  than  mosquitoes. 

Their  life-cycle  has  been  most  completely  worked  out  by  King,  in  Africa. 

The  larvae  and  pupae  occur  in  swiftly  flowing  water,  by  waterfalls,  in  rapids,  etc. 
The  ova  are  laid  in  gelatinous  masses  on  plants  or  rocks  close  to  or  overhanging  the 
water.  The  larva  is  cylindrical,  enlarged  posteriorly,  where  it  is  provided  with  a 
sucker,  by  means  of  which  it  attaches  itself  to  a  rock,  water  weeds,  debris,  etc.  ; 
anteriorly  it  has  a  proleg  close  behind  the  head  on  the  lower  surface.  The  head  is 
dark  and  chitinous.  The  respiration  takes  place  by  means  of  branched  tracheal  gills 
which  protrude  from  the  dorsal  surface  of  the  last  body  segment ;  they  are  retractile. 
The  colour  varies  from  deep  green  to  yellow  or  almost  black.  Their  food  consists  of 
algae  and  other  organisms  in  the  water  brought  to  their  mouth  by  two  fan-like  organs 
.placed  on  the  head.  The  larva?  can  crawl  from  place  to  place  by  means  of  the 
thoracic  proleg ;  they  occur  in  masses,  usually  in  a  more  or  less  erect  attitude.  A  net- 
work of  threads  is  spun  on  their  support,  by  means  of  which  King  tell  us  "  they  are 
enabled  to  maintain  their  position  against  the  strongest  current ;  frequently  they  will 
leave  their  support  and  let  themselves  out  into  the  stream  anchored  by  threads  of 
silk  and  enabled  by  them  to  return." 

When  full  fed  the  larva  spins  a  pocket-shaped  cocoon  on  the  support,  within 
which  it  pupates.  The  pupa  is  motionless  and  has  a  pair  of  branched  spiracles  pro- 
jecting from  behind  the  head.  When  the  adult  emerges,  a  bubble  of  air  collects 
around  it,  and  in  this  it  floats  to  the  surface  and  at  once  takes  wing.  The  European 
species  take  a  month  to  complete  larval  life,  a  week  being  spent  in  the  pupal  stage. 
The  flies  are  most  restless,  and  even  when  stationary  continually  move  their  legs 
about  like  feelers.  Sometimes  the  swarms  consist  entirely  of  females,  sometimes 
early  in  the  season  mostly  of  males. 

The  females  pierce  the  skin  of  humans  on  tender  spots,  such  as  ears,  the  fore- 
head, around  the  eyes  and  nose,  and  crawl  into  the  cavities.  They  are  quite  harmless 
at  night,  mainly  attacking  about  sunrise  and  sunset.  Some  crawl  up  the  arms  and 
legs  and  down  the  neck,  and  leave  behind  little  red  weals  which  itch  intensely 
{S.  dammsum,  Theob.),  and  blood  may  flow  freely  from  the  wounds. 

The  following  are  some  of  the  worst  species : — 

Simulium  columbaschensis,  the  "  Kolumbatz  fly,"  which  abounds  in  the  damp 
marshy  lands  along  the  Danube,  and  is  a  great  plague  toman  and  beasts  in  Hungary, 
and  is  also  abundant  in  Austria  and  Moravia,  and  is  most  numerous  after  inunda- 
tions from  the  Danube.  They  sometimes  appear  in  such  swarms  that  it  is  impossible 
to  breathe  without  getting  them  into  one's  mouth.  There  are  instances  of  children 
being  killed  by  these  flies  when  left  on  the  ground  by  their  mothers  when  working 
in  the  fields. 

S.  damnosum,  Theob.  This  occurs  throughout  Equatorial  Africa  and  is  known 
as  the  «'jinja  fly"  in  Uganda,  the  "fouron"  in  the  French  Congo,  the  "kilteb" 
m  the  Sudan.  It  is  a  most  vicious  biter,  and  in  some  parts  occurs  in  "belts'"; 
Dr.  Christy  found  one  such  extending  from  the  shores  of  the  Victoria  Nyanza 
northwards  along  the  right  bank  of  the  Nile  for  twelve  or  fifteen  miles  or  more,  and 
perhaps  three  or  four  miles  wide.     In  this  area  the  flies  swarm  in  millions  at  certain 


CHIRONOMID^  579 

seasons,  so  much  so  that  the  natives  have  to  leave  their  plantations.  The  bite 
causes  a  weal,  marked  by  a  drop  of  blood. 

S.  gnseicolHs,  Becker.  The  so-called  "  nimitti  "  occurs  in  Upper  Egypt  and 
the  Anglo-Egyptian  Sudan.  It  lives  near  the  river  and  is  not  found  more  than 
half  a  mile  from  it.  Human  beings  are  bitten  on  the  face  and  hands,  animals  in 
the  region  of  the  pudenda. 

S.  lattpes,  Meigen.     This  is  a  European  species,  also  found  in  Natal. 

S.zvellmanni,  Roubaud.  The  "  ohomono  "  of  Angola,  where  it  bites  viciously 
and  is  dreaded  by  the  naked  porters. 

S.  buissoni,  Roubaud.  Occurs  in  abundance  in  the  Marquesas  Islands.  It  has 
been  suggested  that  this  species  may  help  to  propagate  leprosy.' 

A  large  number  of  these  insects  have  been  described  by  Lutz  in  Brazil.-* 

A  Simulium  sp.  (?)  is  very  harmful  to  poultry  in  Cape  Colony.=^ 

In  America,  Simiilidce  are  most  annoying.  One,  S.  meridio)iale,  Riley,  also 
known  as  the  turkey  gnat  in  the  Mississippi  Valley,  has  been  supposed  to  be  the 
carrier  of  chicken  cholera  ;  anyhow,  it  has  caused  the  death  of  thousands  of  chickens 
and  turkeys  in  Virginia  annually.'' 

In  Mexico  Townsend  found  a  Simulium  which  was  named  S.  occideittalis^  which 
caused  great  annoyance  to  man,  many  people  being  so  susceptible  to  them  as  to 


Fig.  401.— Wing  of  Simulium.  Fig.  402.— Wing  of  Chironomus. 

preserve  through  the  gnat  season  a  chronic  inflammation  of  the  exposed  parts  of 
face  and  neck,  resulting  from  the  repeated  bites  giving  rise  to  sores.'* 

Men  and  horses  have  been  partially  incapacitated  by  the  bites  of  sand-flies  or 
Simulium  in  a  Hampshire  wood  (Cantlie,  Brit.  Med.  Jour7i.,  April  28,  1900, 
v.  No.  2,052,  p.   1023). 

Family.    Chlronomidae  (Midges). 

The  Chirotioinidce  or  midges  are  not  only  frequently  mistaken  for  mosquitoes, 
but  some  are  very  annoying  to  man  by  biting  him  as  mosquitoes  do.  They  are 
easily  distinguished  from  true  mosquitoes  {Culicidcs)  by  the  following  characters  : 
(i)  head  small,  often  retracted  under  the  cowl-like  thorax ;  (2)  no  scales  to  the  wings 
or  body;  and  (3)  the  different  arrangement  of  veins  on  the  wings  (fig.  402). 

Two  genera  are  important  as  annoying  man,  namely,  Culicoides,  Latreille,  and 
Johannseniella,  Williston.  The  larvae  of  Chironomidce  are  either  aquatic,  both 
fresh  water  and  marine,  and  help  to  make  the  former  foul,*^  according  to  Slater,  or 
may,  as  in  Ceratopogofiifice,  live  beneath  the  bark  of  trees,  etc.  The  pup£e  are  very 
varied  and  also  the  life-histories  of  the  different  genera.^  The  blood-sucking  habit 
is  confined  to  the  sub-family  Ceratopogofiince . 


1  Btdl.  du  Mus.  d'Hist.  nat,,  1906,  xii,  p.  522. 

^  Mem.  Inst.  Oswaldo  Cruz,  1910,  ii,  fasc.  2,  pp.  211-267. 

'^  C.  Fuller,  "A  New  Poultry  Pest,"  1899,  Leaflet  No.  i,  Dept.  Agric. 

4  Insect  Life,  1888,  i,  p.  14.  ^  Ibid.,  1893,  v,  p.  61. 

"  Entomologist,  1879,  p.  89.       '  Theobald,  "An  Account  of  British  Flies,"  i,  p.  172. 


58o 


THE   ANIMAL   PARASITES   OF   MAN 


Sub-family.   Ceratopogoninae. 


This  sub-family  of  midges  consists  of  very  small  species  varying  from  i  to 
2  mm.  in  length  ;  the  wings  have  darkened  areas,  and  the  second  longitudinal  vein 
is  wanting,  and  the  first  and  third  veins  are  stouter  than  the  others  and  placed  close 
to  the  anterior  margin,  the  fourth  and  fifth  are  forked  ;  the  antennas  in  both  male 
and  female  are  composed  of  fourteen  segments,  six  or  eight  in  the  males  bearing 
long  hairs. 

The  chief  blood-suckmg  species  belong'  to  the  genera  Culicoides,  Latreille, 
and  Johannseniella,  Williston.  The  latter  genus  differs  from  the  former  in  the 
absence  of  an  empodium  or  median  appendage  on  the  last  segment  of  the  tarsi. 
The  genus  Ceratopogon,  as  restricted  by  Kieffer,  is  not  supposed  to  take  vertebrate 
blood,  but  Austen  has  recently  noticed  that  the  type  specimen  of  C.  castafieus, 
Walker,  and  a  new  species  described  by  him,  apparently  have  their  bodies  distended 
with  blood.     The  wings  in  the  Ceraiopo^onincu  are  carried  flat  when  at  rest. 


Fig.  403. — A  Ceratopogon,  or  midge.     Greatly  enlarged. 

In  spite  of  their  small  size  the  females  are  the  most  bloodthirsty  and  annoying  of 
all  insects.  The  Culicoides,  which  are  often  called  "  sand-flies,"  bite  during  the  day 
and  rarely  at  night.  Usually  they  are  most  troublesome  between  3  and  6  p.m. 
They  frequently  attack  in  swarms,  especially  in  the  open,  and  owing  to  their  minute 
size  can  get  through  fine  mosquito  netting.  Some  of  them  produce  a  distinct  "  buzz  " 
when  on  the  wing.  These  insects  are  found  in  all  parts  of  the  world.  No  species 
has  been  definitely  connected  with  any  disease,  but  Culicoides  has  been  suspected  of 
carrying  the  germs  of  Delhi  boil.  The  larvas  of  Culicoides  are  elongate  in  form  and 
have  smooth  bodies  composed  of  thirteen  segments  including  the  head,  which  is 
horny  ;  there  is  no  proleg  on  the  first  segment  as  seen  in  Chironomus,  and  on  the 
anal  segment  are  retractile  gills.  They  are  very  active  and  live  in  the  sap  of  various 
trees  which  saturates  diseased  bark. 

The  pupas  are  smooth,  but  the  abdominal  segments  bear  a  transverse  row  of 
small  spines.  Austen  describes  a  number  of  Culicoides  and  one  Johannseniella  and 
three  Ceratopogons  from  Africa,^  and  Lutz^  a  number  of  this  sub-family  from  Brazil, 


'  Bull.  Ent.  Res.,  1912,  iii,  pp.  99-108. 
-  Mem.  Inst.  Ostvaldo  Cruz,  1913,  v,  fasc. 
pp.  81.99. 


I,  pp.  45-72,  pis.  6-8  ;  and  1914,  vi,  fasc.  2, 


PSYCHODID.^i:  581 

including   a   new   genus,    Centrorhyncus.      Another  genus,    Terseslhes,  Townsend 
(Centrotypus,  Giassi  ;  Mycleiotypus,  Noe),  also  occurs  in  Brazil. 

Culicoides  omatus^  Taylor,  is  described  from  Townsville,  Australia,  found  in 
mangrove  swamps.  It  is  a  very  vicious  biter  and  causes  considerable  irritation, 
settling  on  hands  and  wrists  (Taylor,  Rep.  Ent.  Aust.  Insi.  Trop.  Med.  [191 1], 
191 3,  p.  24). 


Family.     Psychodidae  (Owl  Midges). 

This  family  of  diptera  is  of  considerable  importance,  not  only  on  account  of  the 
blood-sucking  habits  of  some  species,  but  especially  on  account  of  one  at  least  having 
been  proved  to  be  the  carrying  agent  of  "  papataci  "  fever,  a  three-day  fever  very 
prevalent  in  Malta  and  several  parts  of  Southern  Europe  in  the  autumn. 

It  is  also  possible  that  these  small  flies  are  connected  with  the  formation  of 
^'  Delhi  boil,"  caused  by  a  protozoan  parasite. 

PsycJiodidce  are  all  very  small  flies,  many  of  which  have  a  moth-like  appearance, 
and  owing  to  their  fluffy  nature  are  spoken  of  in  Britain  as  "owl  flies,"  sometimes 
also  as  "  window  flies."  Their  bodies  and  wings  are  covered  with  hairs,  densely  in 
some  (sub-family  Psychodino'),  and  in  a  few  with  patches  of  flat  scjuamas.  In  the  non- 
blood-sucking  Psychodiitce  the  wings  are  carried  in  a  peculiar  manner  downwards 
over  the  body,  to  a  slight  extent  resembling  the  Hepialidce,  or  swift  moths.     The 


Fig.  404.— An  owl  midge,  Phlehotomtis  sp.     Greatly  enlarged. 
(From  Giles's  '*  Gnats  or  Mosquitoes.") 

wings  may  be  ovoid  or  lanceolate,  and  have  a  marked  venation  as  seen  in  the  figure. 
The  proboscis  is  short  and  non-suctorial  in  the  majority  of  genera,  but  in  the  sub- 
family Phlebotojuiiice  it  is  elongated  and  hard.  The  antennas  are  long  and  of  sixteen 
segments,  and  bear  whorls  of  fine  hair. 

There  are  two  sub-families,  Psychodincc  2ir\d  Phlebotonii?icc ;  in  the  former  the 
mouth  is  not  suctorial ;  the  female  has  a  horny  ovipositor  and  the  second  longitu- 
dinal vein  is  branched  at  the  root  of  the  wing  ;  in  the  second  sub-family  the  proboscis 
may  be  formed  for  sucking,  the  female  has  no  horny  ovipositor,  and  the  second  long 
vein  has  its  first  fork  near  the  middle  of  the  wing. 

The  sub-family  Phlebotomince  contains  the  genus  Phlebotomus,  which  occurs  in 
South  Europe,  South  Asia,  Africa,  North  and  South  tropical  America.  They  are 
all  small  grey,  brown,  or  dull  yellow-coloured  flies,  and  carry  their  wings  when  at  rest 
upwards  like  a  butterfly.   The  proboscis  is  moderately  long  and  the  legs  long  and  thin. 

The  females  are  most  vicious  blood-suckers,  but  in  some  species  anyhow  the 


582  THE   ANIMAL   PARASITES   OF   MAN 

males  also  bite  {P.  duboscii).  They  are  mainly  nocturnal  feeders  and  hide  away 
during  the  day  in  any  dark  corners  or  crevices. 

The  life-cycle  has  been  worked  out  by  Newstead^  and  Grassi-  in  Europe,  and 
by  Howlett^  in  India. 

The  larv£E  have  been  found  in  crevices  in  rocks  and  caves,  in  dirty  cellars,  and 
dark  damp  places  containing  rubbish,  and  are  also  said  to  live  in  crevices  in  the 
walls  of  privies  and  cesspits. 

The  minute  larva  is  very  marked  ;  as  figured  by  Newstead  it  has  two  long  chaetcC 
projecting  upwards,  in  some  stages  branched,  in  others  simple,  and  on  the  seg- 
ments a  few  blunt  spine-like  processes.  The  pupae  are  found  in  similar  situations. 
The  ova  are  very  minute,  elongate,  translucent  white,  and  covered  with  a  thin  coating 
of  viscous  matter  when  first  laid  ;  soon  after  they  become  dark  brown,  shiny,  with 
long  black  wavy  lines.  Newstead  found  the  incubation  period  in  Malta  to  last  for 
about  nine  days  in  P.papatacii.  Five  species  are  known  in  Europe,  five  in  Africa,^ 
two  in  North  America,  and  eight  are  described  by  Annandale"*  in  the  Oriental  region. 
Lutz  and  Neiva  have  described  three  species  from  Brazil^  {P.  longipalpis^  inter- 
inediiis  and  sqitamiveniris). 

Brachycera  (Flies). 

The  antenna?  as  a  rule  have  three  segments,  and  are  usually  shorter  than  the  head. 
The  first  segment  of  the  antennae  is  frequently  very  small,  and  the  third  one  is  gener- 
ally the  largest,  and  sometimes  possesses  a  terminal  annulated  bristle.  The  palpi  have 
from  one  to  three  segments ;  the  mandibles  are  covered  by  the  labium.  The  three 
thoracic  rings  are  coalesced  ;  wings  are  almost  always  present,  the  posterior  ones 
being  rudimentary  and  covered  with  a  little  scale.  From  the  ova  legless  maggots 
are  hatched,  which  as  a  rule  have  not  a  distinct  head,  but  occasionally  possess 
two  claw-like  booklets.  These  maggots  live  in  decomposing  organic  matter ;  they 
rarely  live  in  water  and  some  of  them  are  parasitic.  They  either  become  barrel- 
shaped  pupas  within  the  last  larval  integument  or,  after  casting  it,  are  trans- 
formed into  naked  pupae.  The  larvae  of  numerous  Brachycera  have  been  observed 
in  man,  some  in  ulcers  or  on  mucous  membranes,  others  in  the  skin  or  in  the 
intestine,  etc.  In  many  cases  the  report  only  mentions  the  presence  of  the  larvae  of 
flies  ;  in  other  cases  the  species  has  been  determined  ;  whilst  in  still  other  cases  the 
corresponding  adult  creature  is  unknown.  We  must  therefore  confine  ourselves  to 
describing  the  most  common  varieties. 

Family.     Phoridae. 

These  flies  belong  to  the  same  division  of  Diptera,  the  Aschiza,  as  the  family 
Syrphidcp  or  "  hover  flies."  They  are  all  small  insects  with  marked  antennae  and 
wings  ;  the  former  have  the  third  segment  globular  and  enlarged,  and  thus  hiding 
the  first  two  ;  the  wings  are  short  and  broad,  the  venation  shows  two  short,  thick, 
long  veins  with  four  thin  ones  running  out  from  them.  The  larvae  normally  live  in 
decaying  animal  and  vegetable  matter,  but  one  species,  Aphiochata  ferruginea^ 
Brun.,  has  been  found  as  an  intestinal  parasite  of  man. 


■  BtiU.  Ent.  Res.,  191 1,  ii,  pt.  2,  pp.  47-78. 

^  "Ricerche   sui   Flebotomi/'  Mem.    delta   Soc.  ital.  delta  Scienze,   1907,     ser.   3,   xiv. 
PP-  353-394- 

»  "  Indian  Sand-flies,"  Ind.  Med.  Cong.,  1909,  sec.  Ill,  pp.  239-242. 

*  Newstead  :  Btttt.  Ent.  Res.,  191 2,  iii,  pp.  361-367. 

^  J^ec.  Ind.  Mus.,  v,  pt.  3,  Nos.  13  and  14. 

"  Mem.  Inst.  Oswatdo  Cruz,  191 2,  iv,  fasc.  I,  pp.  84-95. 


V        SEPSID^,    SYRPHID.E.  583 

Aphiochaeta  ferruginea,  Brun. 

This  small  fly  belonging  to  this  family  is  of  an  orange-ochreous  colour,  the 
upper  part  of  the  thorax  tawny,  and  with  dark  bands  on  the  abdomen,  legs  pale 
yellow,  the  hind  femora  tipped  with  dark  brown.  It  measures  only  2  to  3  mm.  in 
length.  This  insect  is  shown  by  Austen  to  be  widely  distributed  in  the  tropics, 
being  found  in  India,  Burma,  West  Africa,  and  Central  America.  The  larvae  breed 
in  decaying  animal  matter,  such  as  putrid  meat,  decomposing  shell-fish,  etc. 

Heusner  bred  out  sixty-three  flies  from  larvae  taken  from  an  Indian's  foot. 

Baker  {Proc.  Binnia  Brafich  Brit.  Med.  Assoc,  1891,  p.  11- 16)  found  that 
the  maggots  of  this  fly  were  passed  per  aiium  by  a  European  at  intervals  during 
a  period  of  ten  months.  Baker  found  that  the  larvre  fed  on  human  faeces;  from 
the  egg  stage  to  the  deposition  of  eggs  from  the  resultant  brood  of  flies  occupied 
twenty-two  days.  He  concludes  that  they  are  capable  of  propagating,  and  do 
so  while  living  within  the  human  intestines.  He  also  records  the  larvae  in  two 
girls. 

The  larva  does  not  seem  to  have  been  described,  but  Austen  describes  the  pupa 
{Trans.  Soc.  Trap.  Med.  and Hyg.^  iii.  No.  5,  p.  229). 

Phora  rufipes,  Meig. 

The  larvie  of  the  "hump-backed  fly"  live  in  rotting  potatoes,  muslirooms, 
radishes,  etc.,  and  when  accidentally  introduced  into  the  intestine  of  man  can, 
like  other  larvae,  live  there  twenty-four  hours  and  even  more,  and  may  set  up  serious 
gastric  disturbances. 

P.  rufipes  is  the  same  as  P.  pallipes.,  Latr. 

Family.     Sepsidae. 

Small  blackish  flies,  elongate,  with  abdomen  narrowed  at  the  base,  thickened 
and  curved  downwards  towards  the  extremity.  Larvae  often  found  in  decaying 
vegetables,  ham,  cheese,  etc.  The  larvae  have  the  power  of  skipping  ;  conical  in 
form,  pointed  in  front,  truncated  behind,  about  5  mm.  long,  shiny  and  smooth,  the 
anal  segment  with  fleshy  protuberances.  The  genus  Piophila  has  a  short  proboscis 
and  the  cross-veins  of  the  wings  approximate. 

Piophila  casei,  L. 

Cheese  flies.  The  larvae  live  in  ripe  cheese,  with  which  they  are  sometimes 
introduced  into  human  beings  (Meschede). 

The  larvae  of  the  cheese  flies  {Piophila  casei)  may  pass  through  the  alimentary 
canal  of  human  beings  alive,  and  have  been  occasionally  referred  to  in  cases  of 
internal  myiasis.  It  also  breeds  in  dead  bodies  in  adipose  tissue.  Howard  records 
it  on  human  excrement.  It  is  thus  possible  that  some  of  the  recorded  cases  of  thi^ 
pest  being  passed  alive  may  be  due  to  eggs  deposited  on  human  faeces. 

Family.     Syrphidae  (Hover  and  Drone  Flies). 

Amongst  the  large  family  of  Syrphidce  is  found  a  section  known  as  the  Eristalince 
or  drone  flies,  whose  curious  long-tailed  larvae  are  popularly  called  "  rat-tail 
larvae,"  on  account  of  the  end  of  the  body  being  drawn  out  into  a  long  telescopic  tail 
of  two  segments,  at  the  end  of  which  are  placed  the  breathing  pores.     These  larvae 


384  THE   ANIMAL   PARASITES   OF    MAN 

live  in  water,  no  matter  how  foul,  and  in  liquid  manure.  They  have  occasionally 
been  obtained  in  foul  drinking  water  by  human  beings  and  from  eating  watercress 
improperly  washed  or  from  badly  kept  beds.  Austen  {Trans.  Soc.  Trop.  Med.  and 
Hyg.,  iii,  No.  6,  p.  221)  records  that  in  the  autumn  of  1907  a  number  of  the  larvae 
of  the  common  drone  fly  {Eristalis  tenax)  were  passed  per  reciiun  by  a  woman  in 
Hampshire  who  had  recently  arrived  from  France.  The  patient  had  eaten  a  con- 
siderable quantity  of  watercress  before  leaving  France.  I  have  twice  found  small 
Eristalis  larva?  clinging  by  their  long  tails  on  watercress  served  at  table. 

Family.     Drosophilidae. 

Small,  rather  plump  flies,  with  short,  broad  abdomen,  with  bristles  on  the  head 
and  legs.     Often  abundant  in  decomposing  fruit,  and  may  occur  in  dense  masses. 

Drosophila  melanogaster,  Br. 

The  larva?  of  this  fly  occur  in  over-ripe  fruit  and  in  fungi,  often  also  in  human 
habitations,  and  live  in  substances  undergoing  acid  fermentation  (vinegar,  decaying 
fungi,  rotting  fruit,  in  damaged  spots  in  diseased  trees),  much  more  rarely  in  animal 
substances,  and  they  occasionally  gain  access  to  the  human  intestine  (for  example,  by 
the  medium  of  sour  milk).  When  introduced  in  any  quantity,  they  cause  vomiting 
or  attacks  resembling  colic  ;  when  taken  in  the  pupal  stage  no  unpleasant  results 
are  produced. 

Family.     Muscidae. 
Teichomyza  fusca,  Macq. 

Syn.  :  Scatella  in'inaria.,  Rob.  Desv.  ;  EpJiydra  loncripetinis.,  Meigen. 

The  larvae  live  in  the  urine  in  privies.  Several  authors  state  they  have  found 
them  in  fresh  fasces  or  in  vomited  matter.  Pruvot  states  that  they  continue  for  three 
days  in  the  stomach  of  rats  into  which  they  have  been  intentionally  introduced. 
(Pruvot,  G.,  "  Contrib.  k  I'etude  des  larves  de  dipt,  trouv.  dans  le  corps  humain," 
These  de  Par..,  1882;  Chatin,  J.,  in  Coinp.  rend.  Soc.  de  BioL.,  Paris,  1888  [8],  v, 
p.  396  ;  Roger,  H.,  ibid..,  1851  [i],  iii,  pp.  88,  etc.) 

Homalomyia   canicularis,   L.,  etc. 

Hoiiialoniyia  nianicata,  Meig.,  live  as  larvc-e  in  decomposing  vegetable  matter  or 
in  cultivated  vegetables  (cabbage)  ;  they  are  easily  recognizable  by  their  plumed 
bristles,  which  are  situated  laterally  on  the  body  segments.  They  obtain  access 
fairly  often  to  the  human  intestine  and  give  rise  to  very  uncomfortable  symptoms. 
Cases  have  been  recorded  from  Germany,  Austria,  France,  England,  North 
America  (Wacker,  in  Artzl.  Intelligenzbl.,  1883,  xxx,  p.  109;  Florentin,  in  Compt. 
rend.  Soc.  de  Biol,  Paris,  1904,  Ivi,  p.  525;  and  other  authors). 

The  larvae  of  an  allied  genus  (Anthomyia),  which,  however,  does  not  possess 
plumed  bristles,  has  been  found  in  the  external  auditory  meatus  of  a  man  {A.plicvialis, 
according  to  Danthon). 

[//.  cafiicularis  is  common  to  Europe  and  North  America,  and  is  an;abundant 
house-fly.  It  is  the  small  house-fly  so  often  seen  on  windows.  Besides  living  on 
vegetable  matter,  they  have  also  been  found  in  the  nests  of  the  humble  bee.  Larvae 
of  this  species  (fig.  405)  were  sent  to  the  British  Museum,  taken  from  the  f^ces  o 


MUSCID^ 


585 


a  woman  suffering  from  cancer.'     They  were  found  at  Shrewsbury.     Hagen^  reports 
the  larvc^  of  this  fly  as  occurring  alive  in  the  urethra  of  a  patient. — F.  V.  T.] 

Homalomyia  scalaris,    Fabr. 

[This  is  not  a  synonym  of  the  above,  but  a  distinct  species. 
[//.  )iianicata^  Meigen,  is  also  distinct.— P".  V.  T.] 


Anthomyia    desjardensii,   Macq. 

This  fly,  allied  to  Homaloinyia,  is  the  cause  of  both  intestinal  and  cutaneous 
myiasis  at  Bihe,  Angola  (Wellman,  Joiirn.  Trap.  Med.  and  Hyg.,  June,  1907, 
X,  p.   186). 

Hydrotaea    meteorica,   L. 

The  larvae  live  in  decaying  vegetable  substances,  also  in  dung,  and  have  been 
evacuated  in  some  cases  by  man  (Zetterstedt,  Joseph). 


Fig.  405. — Larva  of 
Homalomyia  canictilaris. 
Enlarged. 


Fig.  406. — Larvse  of 
Calliphora  vojuitoria. 
Enlarged. 


Fig.  407.  —  Larva  of 
Chrysomyia  nmcellaria. 
4/1.     (After  Conil.) 


Cyrtoneura   stabulans. 

Larvae  in  fungi,  but  occasionally  also  on  larvae  of  butterflies  and  Hynienoptera_ 
occasionally  introduced  into  the  human  intestine  (Joseph). 


Musca  domestica,   L., 

and  M.   {Calliphora)  vomitoria^   L.,   and  allied  species  ;  larvae  of  these  have  been 
repeatedly  found  in  the  intestine  and  nose  of  man  (Mankiewicz,  etc.).* 

'  Theobald,  "  First  Report  Economic  Zoology,"  Brit.  3fus.  {Nat.  Hist.),  p.  55. 

'^  Hagen,  Froc.  Bost.  Soc,  N.H..,  xx,  p.  107. 

^  "  Larvae  of  a  Musca,  probably  M.  corvina,  were  passed  in  numbers/^;-  rectiim  by  a  child 
in  Liverpool  with  Homalomyia  larvre," — "Second  Report  Economic  Zoology,"  Theobald, 
1903,  p.   16. 


586 


THE   ANIMAL   PARASITES   OF    MAN 


Musca  domestica,  Linn.  (Common   House-fly) 


It  is  not  only  on  account  of  a  few  larvae  of  the  common  house-fly  {Musca 
domestica)  being  found  in  the  intestines  of  man  that  it  is  of  importance  medically. 
It  is  far  more  important  on  account  of  the  part  it  plays  in  the  spread  of  diseases 
of  the  intestines,  such  as  typhoid  fever  and  cholera,  infantile  diarrhoea  and 
dysentery. 

Howard  and  Clark  {Journ.  Exp.  Med.,  1912,  xvi.  No.  6,  pp.  850-859)  have  shown 
that  the  house-fly  is  capable  of  carrying  the  virus  of  poliomyelitis  for  several  days 
on  the  surface  of  the  body  and  for  several  hours  in  the  gastro-intestinal  tract. 
The  house-fly  may  also  distribute  the  ova  of  Tccfiia  solium  and  the  white  worms 
(Oxyuris  and  Ascaris).  It  has  also  been  proved  that  they  may  carry  the  germs 
of  tuberculosis,  and  it  is  said  that  they  play  an  important  part  in  the  spread 
of  infectious  ophthalmia  in  Egypt. 

This  insect  is  found  in  all  parts  of  the  world.  In  warm  countries  it  breeds 
all  the  year  round,  and  it  may  do  so  even  in  temperate  climates  in  warm  places, 
such  as  stove  houses.  Most,  however,  die  off  in  the  autumn  ;  but  some  survive  the 
winter  as  adults,  in  such  places  as  kitchens,  restaurants,  and  warm  houses.  I  have 
never  failed  to  find  a  few  Musca  domestica  in  houses  during  the  winter.  The 
majority,  however,  hibernate  as  puparia. 

The  females  deposit  from  120  to  150  eggs  in  a  batch  in  stable  manure,  rotting 
vegetation,  house  refuse,  spent  hops,  old  soiled  bedding,  etc.  A  single  female 
may  lay  as  many  as  six  batches  of  ova  during  her  life.  The  eggs  are  shiny  white, 
and  hatch  in  from  eight  to  twenty-four  hours  in  warm  weather  to  three  or  four 
days  in  cool  weather.  The  while  footless  maggots  are  cylindrical,  tapering  to 
a  point  at  the  head  end,  truncated  posteriorly.  The  head  consists  of  two  dark 
mandibular  hooks  and  two  short  antennse.  ,On  the  tail  end  are  two  plates,  the 
stigmata,  in  which  the  main  tracheal  trunks  open  ;  in  the  second  segment  are 
a  small  pair  of  projecting  stigmata.  The  larval  stage  lasts  from  seven  to  five 
days  in  hot  weather  ;  but  in  cold  weather  in  temperate  climes  it  may  last  six  or 
eight  weeks. 

The  larva  on  reaching  maturity  becomes  a  barrel-shaped  puparium  of  a  dark 
brown  to  black  colour,  and  in  this  case  changes  to  the  pupa.  This  stage  lasts  from 
three  days  in  the  tropics  to  four  or  five  weeks  in  cold  weather,  the  life- cycle  thus 
varying  from  ten  days  in  the  tropics  io  fourteen  in  warm  weather  in  Europe  up  to 
three  or  four  months  under  unfavourable  conditions. 

All  breeding  grounds  should  be  burnt  or  otherwise  done  away  with,  such 
as  stable  manure,  house  and  kitchen  refuse,  human  excrement  and  soiled  substances, 
also  decaying  vegetation  as  soon  as  possible,  certainly  by  every  sixth  day.  Stable 
manure  should  be  kept  in  closed  receptacles  and  should  be  removed  by  every  sixth 
day  to  at  least  one  mile  from  habitations  and  sprinkled  with  chloride  of  lime. 
All  kitchen  and  household  refuse  should  be  burnt  at  once  or  buried  in  pits  and 
covered  with  soil.  Latrines  should  be  as  far  as  possible  from  hospitals,  mess  rooms 
and  tents.  Food — especially  milk,  sugar  and  fruit — should  be  kept  screened  with 
muslin  when  house-flies  are  about.  Mess  rooms  and  tents  and  hospitals  should 
have  doors  and  windows  screened  with  fine  wire  gauze  during  the  fly  season. 
All  possible  steps  should  be  taken  to  prevent  them  contaminating  man's  food  and 
from  breeding  in  human  excrement  and  from  entering  hospitals.  When  present 
in  dwelling-houses  in  numbers  they  may  be  killed  by  fumigation  with  pyrethrum 
or  sulphur. 


CHRYSOMYIA 


587 


Genus.     Chrysomyia,  Rob.  Desv. 

Chrysomyla   (Compsomyla)    macellaria,    Fabr. ; 
Lucilia  macellaria,  Fabr. 

Syn. :  Lucilia  /lofninivorax,  Coq. ;  Calliphora  itifesta,  Phil.  ;  Calliphora 
anthropophaga,  Conil. 

A  species  distributed  from  the  Argentine  to  the  south  of  the  United  States 
which  deposits  its  ova  on  ulcers,  in  the  aural  meatus  or  in  the  nasal  cavities  of 
persons  who  sleep  in  the  open  air.  The  larvae  are  yellowish  white,  16  mm.  long,  are 
armed  with  two  strong  mouth  hooks,  and  provided  with  spinous  rhigs  (screw-worm)  ; 
they  lie  hid  in  the  nasal  and  frontal  sinuses,  in  the  pharynx,  larynx,  etc. ;  they  per- 
forate the  mucOus  membranes,  even  cartilage,  migrate  into  the  eyes,  the  cranial 
cavity,  middle  ear,  and  cause  severe  disturbances  ;  after  the  mature  stage,  in  which 
the  larvae  leave  the  host  to  enter  the  pupal  state,  these  symptoms  often  spontaneously 


Fig.  408. — The  screw-worm  fly  {Chrysomyia  niacellaria), 

abate  after  a  lapse  of  eight  days,  leaving  behind  greater  or  less  cicatrices,  and  con-^ 
sequently  also  defects  in  function  of  the  organs  attacked.  Very  often,  however,, 
sepsis  sets  in,  usually  with  a  fatal  termination. 

(Coquerel  in:  Arch.  gen.  de  med.,  1858  (5),  p.  513;  1859,  xiii,  p.  685;  Ann.  Soc.  enl. 
France^  1858  (3),  vi,  p.  171  ;  1859,  vii,  p.  234.  Weber  in  :  Rec.  de  mem.  de  mtd.  milit.,  1867 
(3),  xviii,  p.  159.  Franclus,  A.,  in:  Arch.  f.  path.  Anat.,  1868,  xliii,  p.  98.  Conil  in  : 
Bol.  Acad.  nac.  cienc.  Cordoba^  i88r,  iii,  p.  296.  Humbert,  Fr.,  in  :  Proc.  U.S.  Nat.-  Mns.,. 
1883,  vi,  p.  103;  Amer.  Nat.,  1884,  xviii,  p.  540.  Lindsay  in  :  Journ.  Trap.  Afed.,  1902; 
v,  p.  220,  and  other  authors.) 

[This  species  is  known  as  the  screw-worm  fly.     It  attacks  animals  as  well  as. 
man,  especially  laying  its  eggs  on  wounds  formed  by  barbed  wire.     It  may  also  be 
found  on  dead  flesh.     Dr.  St.  George  Gray  sent  me  speciniens  from  St.  Lucia,  from 
37 


588  THE   ANIMAL   PARASITES   OF   MAN 

the  nose  and  mouth  of  a  patient  in  Victoria  Hospital.  Others  were  found  in  the 
vagina  of  another  patient.  Out  of  the  four  patients  attacked,  two  occupied  the 
same  bed,  one  after  the  other,  and  a  third  the  next  bed  to  it.  The  other  case  was 
in  a  more  remote  part  of  the  hospital.  There  are  numerous  records  of  this  fly 
attacking  man.     It  occurs  from  the  Argentine  to  Texas. — F.  V.  T.] 

Chrysomyia  viridula,  Rob.  Desv. 

[This  species  is  somewhat  larger  than  the  former ;  the  body  is  metallic  bluish- 
green,  the  dorsum  of  the  thorax  with  three  blackish,  longitudinal  stripes,  and  the 
face  ochraceous ;  about  lo  mm.  long.  Austen  records  this  species  from  man, 
Dr.  Daniels  having  bred  it  from  larvae  from  a  sore  on  a  human  being  in  New 
Amsterdam,  British  Guiana.  Dr.  Laurence  also  bred  it  in  Trinidad.  In  the  latter 
case  between  loo  and  150  maggots  were  discharged  from  the  nose  of  a  woman 
suftering  from  facial  myiasis  (^Brit,  Med.  Joicrn.,  January  9,  1909,  p.  88  +  fig.). 
— F.  V.  T.] 

Genus.     Lucilia,  Rob.  Desv. 

Lucilla  nobilis,  Meig. 

The  larvae  were  observed  by  Meinert  in  Copenhagen  in  the  auditory  meatus 
of  a  person  who,  after  taking  a  bath,  fell  asleep  in  the  open  air,  and  on  waking 
felt  singing  in  the  ears,  and  had  a  sensation  as  if  there  were  water  in  the  auditory 
canal.  During  the  next  days  severe  pains  set  in,  and  there  was  a  discharge  of 
blood  and  pus  from  both  ears,  as  well  as  from  the  nose.  On  washing  out  the 
meatus  the  maggots  made  their  appearance. 

Lucilia  ccesar  and  Z.  sericata  have  also  been  observed  in  the  larval  state  in 
man  (Thompson,  Hope,  Henneberg  and  Calendoli,  Napoli,  1907). 

[This  golden-green  fly  usually  lays  its  eggs  on  decomposing  organic  matter ; 
now  and  again  it  lays  its  eggs  in  wounds  on  man. — F.  V.  T.] 

Genus.     Pycnosoma,  Brauer  and  v.  Bergenstamm. 

The  species  of  this  genus  have  a  general  resemblance  to  the  Lucilias  and 
Chrysomyias,  but  the  body  is  stouter  and  the  abdomen  banded.  The  genus  can  be 
distinguished  from  Chrysomyia  by  the  absence  of  the  three  thoracic  stripes  and 
by  the  eyes  of  the  male,  in  which  the  facets  forming  the  upper  portion  are  much 
enlarged,  whereas  in  Chrysomyia  they  are  not  noticeably  larger.  Austen  also  points 
out  that  the  sterno-pleural  bristles  in  Pycnosoma  are  i  :  i,  in  Chrysomyia  2:1. 
The  genus  is  found  in  tropical  Asia  and  Africa  only.  All  records  of  Chrysomyia 
(Compsomyia)  in  India  must  be  referred  to  this  genus.  Bezzi  and  Stein  ("  Katalog 
der  Palaarktischen  Dipteren,"  1907,  iii,  p.  543),  however,  regard  the  two  as 
synonymous. 

The  larvae  are  frequently  found  in  the  nostrils  of  man  and  burrow  into  the  sinus, 
but  normally  they  live  on  decaying  animal  matter. 

Pycnosoma  forms  the  so-called  Indian  screw-worm.  Patterson  {Ind.  Med.  Gaz.^ 
October,  1909,  xliv,  No.  10)  records  the  case  of  a  woman  at  Tezpin,  Assam,  from 
whom  as  many  as  100  larvae  were  removed  at  one  time,  and  later  the  left  orbital 
cavity  was  found  packed  with  hundreds  of  maggots;  eventually  the  patient  died. 
It  is  possible  that  this,  however,  was  due  to  a  species  of  Sarcophaga.  Austen 
undoubtedly  records  this  genus  causing  nasal  myiasis  in  India  {Trans.  Soc.  Trop. 
Med.  and  Hyg.,  iii,  p.  235).  At  Dehra  Doon,  U.P.,  a  woman  discharged  100 
larvae  from  her  nose,  with  great  pain  in  the  nasal  region  and  frontal  sinuses. 

The  so-called  "  peenash,"  a  common  malady  in  Rajputana,  is  a  true  nasal  myiasis. 


SARCOPHAGA  589 

Genus.     Sarcophaga,   Mg. 
Sarcophaga  carnosa,  L.,  1758. 

Larvae  of  flesh-flies  provided  with  two  claws  at  the  anterior  end,  which  settle 
on  raw  or  cooked  meat,  and  in  the  open  on  carcases  of  animals  ;  they  are  often 
observed  in  man,  both  in  the  intestine  (introduced  with  food)  and  in  the  nasal 
cavities,  frontal  sinus,  conjunctiva,  aural  meatus,  anus,  vulva,  vagina,  prepuce, 
and  open  ulcers,  often  migrating  further  from  the  regions  first  attacked.  (Gayot 
in  Compt.  rend.  Acad.  Set.,  Paris,  1838,  vii,  p.  125.  Grube  in  Arch.  f.  Naiurg.,  1853, 
xix,  I,  p.  282.  Legrand  du  Saulle  in  Compt.  rend.  Acad.  Set.,  Paris,  1857,  xlv,  p.  600, 
and  other  authors.) 

[This  fly  is  viviparous.  The  fly  varies  from  10  to  30  mm.  in  length,  and  is 
of  a  general  ash-grey  colour ;  the  thorax  with  three  dark  stripes,  the  abdomen 
light  grey  with  three  black  spots  on  each  segment  ;  legs  black  ;  base  of  wing.s 
yellow.  It  also  attacks  animals  and  birds,  especially  geese.  The  genus  Sarcophaga 
is  universally  distributed.  The  maggots  are  whitish  or  yellowish  footless  larvas 
of  twelve  segments,  tapering  to  a  point  in  front,  broadened  posteriorly.  There 
are  two  mouth  liooks,  by  means  of  which  they  rasp  their  food.  The  breathing 
pores  are  at  the  end  and  consist  of  two  groups  of  three  slits,  each  surrounded  by 
a  hardened  area.     They  pupate  in  their  old  skin,  which  turns  brown. —  F.  V.  T.] 

Sarcophaga  magnifica,  Schiner,  1862.^ 

Syn.  :  Sarcophaga  wohlfahrti.,   Portschinsky,   1875. 

A  species  widely  distributed  over  the  whole  of  Europe,  occurring  especially  in 
Russia  (Mohilew)  ;  the  presence  of  the  larvse  in  man  was  first  observed  by 
Wohlfahrt  (1768).  The  larvae  settle  in  the  pharynx,  in  the  nose,  the  aural  meatus, 
the  conjunctiva,  and  in  other  regions  of  the  human  body  ;  they  also  attack  domestic 
animals  and  birds.  As  Portschinsky  has  shown,  they  cause  severe  inflammations, 
haemorrhages  and  suppurations  in  the  organs  in  which  they  occur  ;  children  are 
especially  attacked.  A  number  of  cases  have  been  observed  also  in  Central  and 
Western  Europe.  [The  fly  has  a  light  grey  abdomen  with  shiny  black  spots  which 
do  not  change  their  shape  and  appearance  according  to  the  angle  in  which  the  fly 
is  viewed. — F.  V.  T.] 

(Wohlfahrt :  "  Observ.  de  vermibus  per  nares  excretis,"  Halae,  1768  ;  Nov.  Act. 
Acad.  Caes.  Nat.  curios..,  1770,  iv,  p.  277.  Gerstacker  in:  Sitzicngsb.  Ges.  nat.  Frde. 
Berl.,  1875,  p.  108.  Portschinsky  in:  Horce  soc.  efttom.  ross.,  1875,  1884,  p.  123. 
Laboulbene  in  :  Ann.  Soc.  ent.  France,  1883  (6),  iii  ;  Bull.,  p.  xcii.  Leon  in  :  Bull. 
Soc.  des  MM.  et  Nat.  de  /assy,  1905,  xix,  p.  i.  Freund,  L.,  in  :  Verh.  Ges.  deutsch. 
Naturf.  u.  Arzte,  Homburg  [1901],  1902,  ii,  2,  p.  450,  and  other  authors.)  [Probably 
most  cases  of  attack  in  Europe  are  due  to  this  species. — F.  V.  T.] 

The  above  cited  do  not  exhaust  the  number  of  observations  of  diptera  larv;e 
parasitic  in  man  ;  there  are  yet  to  be  mentioned  the  larvae  of  .S".  hcemorrhoidalis , 
S.  hcEinatodes  (of  G.  Joseph),  those  of  S.  ruficornis  (excitants  of  a  cutaneous  myiasis 
in  the  East  Indies),  those  of  species  of  Eristalis  (of  Hanby  and  others),  and  those  of 
Fhora  rufipes  (of  Kahl,  of  Warsaw,  and  others).  In  many  cases  the  determination 
of  the  diptera  laivaj  has  been  omitted  (or  must  be  omitted) ;  such  is  the  case  with 
diptera  larvae  in  the  eye  (Schultz-Zehden  in  :  Berl.  klin.  Wochenschr.,  1906,  p.  286. 
OUendorf  in  :  Med.  Korrespo7idenzbl.  d.  wiirt.  drtzl.  Landesver.,  1904,  p.  1017. 
Kayser  in  :  Kli7t.  Monatsbl.  f.  Augenheilkunde,  1905,  xliii,  i,  p.  205.  Ewetzky  and 
V.  Kennel  in  :  Zeitschr.f.  Augenheilktuide,  1904,  xii,  p.  337,  and  other  cases).    Austen 

'   [The  correct  name  for  this  fly  is  Wohlfahrtia  magnifica,  Schiner. — F.  V.  T.] 


590 


THE   ANIMAL   PARASITES   OF   MAN 


records  several  cases  of  myiasis  due  to  Sarcophaga  {vide   Trajis.  Soc.  Trap.  Med. 
and  Hyg.,  1910,  iii,  No.  6). 

The  larvae  of  African  Muse  Ida'  have  now  become  of  greatei- 
interest;  Hke  several  Oestrid  larvae  they  live  normally  in  the  skin  of 
mammals,  but  also  attack  man.  The  knowledge  of  these  species  is 
certainly  very  insufficient,  but  this  is  not  likely  to  be  the  case  much 
longer,  as  medical  men  practising  in  the  Colonies  are  giving  their 
attention  to  these  parasites.  At  the  present  time  four  distinct  forms 
are  recognized  according  to  Gedoelst.^ 

Sarcophaga  chrysdtoma,  Wied. 

[This  species  is  recorded  as  attacking  human  beings  at  New  Amsterdam,  Britisli 
Guiana.  The  fly  is  15  mm.  long,  has  a  golden-coloured  face,  three  broad  black 
thoracic  stripes  and  ochraceous  buflf  anal  segments.  It  was  bred  from  larvife 
obtained  by  Dr.  Roland  from  a  sore  on  a  girl's  foot.  It  is  known  to  occur  in  the 
Brazils  and  the  West  Indies.  Another  species  was  also  bred  which  Austen  was 
unable  to  identify. — F.  V.  T,] 

Sarcophaga  plinthopyga,  Wied. 

[This  and  other  species  of  Sarcophaga  are  called  "yaw  flies"  in  Dominica,  as 
they  are  believed    to  be    concerned  in  the   dissemination    of  frambcesia    or   yaws 
(Nicholls)  {vide  Austen,  Trans.  Soc.   Trop.  Med.  and  Hyg.^ 
1910,  iii,  p.  239).— F.  V.  T.] 

Ochronnyla  anthropophaga,  E.  Blanch. ; 
Cordylobia  arthrophaga,  Griinberg. 

Indigenous  to  the  Senegal  and  neighbouring  districts  ; 
in  the  district  of  Cayor  (between  the  mouth  of  the  Senegal 
and  Cape  Verde)  the  larva  is  known  as  the  "  ver  de  Cayor."' 
It  lives  under  the  skin,  especially  at  the  lower  extremities 
and  the  lower  region  of  the  trunk,  producing  small  boils, 
which  cause  pain,  but  after  about  eight  days,  when  the 
larva  leaves  the  body  to  enter  the  pupal  stage,  the  pain 
discontinues.  Besides  man  the  larva  occurs  in  dogs,  goats, 
cats,  and  in  the  jackal.  It  is  still  questionable  whether 
the  fly  deposits  its  eggs  direct  or  on  the  ground,  from 
whence  the  larvae  as  they  emerge  gain  access  to  animals 
and  man.  Larvae  yellowish-white,  14  mm.  long,  4  mm.  wide, 
eleven  segments^  ;  head  with  two  globular  antennas-like 
appendages,  two  black  curved  mouth  hooks,  and  two  wart- 
shaped,  finely  spinous  structures  at  their  base.  Body  evenly 
covered  to  the  seventh  segment  with  small  black  prickles, 
which  are  stronger  at  the  sides  and  the  anterior  borders  of  the  segments ;  from  the 
seventh  they  increase  in  size,  on  the  two  hindermost  they  are  wanting ;  on  the  last 
segment  two  deep  yellow  spiracles,  each  with  three  markedly  curved  fissures ;  in 

'  [The  following  are  known  to  cause  myiasis  in  man  in  Africa  :  Cordylobia  anthropophaga, 
Griinb.  ;  Aiichmeromyia  liiteola,  Fabr.  ;"  A.  rodhani,  Gedoelst  ;  Oestrus  ovis,  Linn.  ;  and 
Anthomyia  desjardensii,  Macq.  The  anthropophaga,  Blanchard,  and  the  depressa,  Walker, 
referred  to  here  are  Griinberg's  anthropophaga. — F.  V.  T.] 

2  [Austen  gives  the  length  as  12  to  12-5  mm.  and  the  breadth  as  5  mm.  ;  he  describes  the 
larva  as  follows  :  Bluntly  pointed  at  the  anterior  extremity,  and  truncate  behind  ;  from  third 


Fig.  409. — Ochromyia 
larva  on  the  skin  of 
man,  South  Africa. 
3/1.     (After  Blanchard.) 


CORDYLOBIA  59 1 

addition  two  stigmata  on  the  posterior  border  of  the  first  segment.  Duration  of  the 
larval  stage  about  eight  days.  Upon  the  construction  of  roads  in  Guinea  the  larva 
is  spread  by  dogs  far  into  the  interior. 

Auchmeromyla  (Bengalia)  depressa  (Walker).^ 

Distributed  in  the  region  of  Natal  and  apparently  over  the  whole  of  South 
Africa.  The  "  larva  of  Natal,"  as  one  may  still  term  the  species  provisionally,  as 
its  identity  is  not  certain,  possesses  on  its  head  (besides  the  mouth  hooks)  lateral 
protuberances  beset  with  a  row  of  chitinous  spines.  The  cuticle  of  the  body  is 
spinose.  The  spines  are  difficult  to  recognize  on  account  of  their  transparency  and 
want  of  colour;  they  are  longest  over  the  anterior  segments,  from  the  fifth  they 
become  smaller,  and  over  the 
hindermost  they  are  very  small. 
Apart  from  the  foremost  segment, 
the  position  they  take  is  that 
of  rows  running  transversely  or 
obliquely,  two  to  four  generally 
in  juxtaposition  ;  the  number  of 
spines  in  the  groups  gradually 
increases  posteriorly,  attaining  the 
number  of  eight  to  twelve  on  the 
sixth  segment,  and  this  number  is 
maintained  to  the  end  of  the  body. 
Isolated  spines  are  found  over  the 
head  ;  over  the  second,  third  and 
fourth    segments    single    ones   are 

still  found  adjoining  the  groups  of  -n^  tj~~^      j    c  ui  r  -nt  .  . 

r  \       r  °        ^  Fig.  410. —Head  end  of  "  larva  of  Natal, 

spmes,  from  the  filth  onward  they  Magnified.     (After  Gedoelst.) 

are  wantmg.    From  here  the  spines 

cover  the  whole  free  surface  of  the  segments  ;  over  the  fourth  the  anterior  three- 
quarters,  over  the  third  two-thirds  and  over  the  first  and  second  only  the  anterior 
half.  The  stigmata  found  at  the  anterior  end  also  serve  as  distinguishing  characters. 
The  parasitic  stage  appears  to  last  about  fourteen  days.  [Fuller  {Agric.  Joiirn.^ 
Dept.  Agric.  and  Mines,  Natal,  1901,  iv,  p.  606)  refers  to  this  as  Bengalia  depressa 
also.  — F.  V.  T.] 

Genus.     Cordylobia,  Griinberg,  1903. 
Cordylobia  grlinbergi,  Donitz. 

Syn.  :   Ochroniyia  anthropophaga^  Griinberg,  7iec  Blanch. ;   Cordylobia 

ajithropophaga^  Griinberg. 

Endemic   in    German    East   Africa   and  neighbouring  regions.      Larva   up   to 

14  mm.  long,  4  to  5-5  mm.  wide,  of  cylindrical  shape,  slightly  narrowed   behind, 

truncated,  gradually  tapering  in  front ;   antenna-like  processes,  cone-shaped,  blunt. 

to  eleventh  segments  thickly  covered  with  minute  recurved  spines  of  brownish  chitin,  usually 
arranged  in  transverse  series  of  groups  of  two  or  more,  which  can  be  seen  to  form  more  or  less 
distinct  undulating  and  irregular  transverse  rows.  In  each  of  the  two  posterior  sligmatic 
plates,  the  respiratory  slit  on  either  side  of  the  median  one  is  characteristically  curved, 
resembling  an  inverted  note  of  interrogation.  The  barrel-shaped  puparium  is  on  an  average 
iO'3  by  4*6  mm.  ;  its  colour  varies  from  ferruginous  to  nearly  black. — F.  V.  T.] 

'  [According   to   Austen  this   is   Cordylobia  anthropophaga^  Griinb.     Bengalia   depressa^ 
Walker,  is  a  very  different  insect,  whose  life-history  is  unknown. — F.  V.  T.] 


^g2  THE   ANIMAL   PARASITES   OF   MAN 

Smaller  cylindrical  formations  at  the  base  of  the  mouth  hooks  surrounded  by  a 
circle  of  chitinous  hooks.  Body  from  the  first  segment  covered  with  small  brown 
squamous  spines  which  are  disposed  in  numerous  irregular  transverse  rows.  The 
spines  are  small  over  the  two  first  segments,  the  two  posterior  thirds  of  all  the 
segments,  as  well  as  from  the  eighth  ;  over  the  third  to  the  seventh  they  are  larger, 
but  between  these  there  are  very  small  spines.  The  breathing  pores  of  the  stigmata 
at  the  anterior  end  are  kidney-shaped  ;  the  orifices  are  elongated  and  very  tortuous, 
each  divided  into  three.     The  larval  period  appears  to  last  several  weeks. 

Cordylobia  anthropophaga,  Grunberg. 

This  well-known  cutaneous  African  parasite  seems  to  have  been  the  cause 
of  much  confusion  in  regard  to  names.  It  belongs  to  the  genus  Cordylobia 
of  Grunberg,  and  is  one  of  the  family  Muscidcp,  and  differs  from  Auchmeromyia 
in  that  the  second  abdominal  segment  of  the  female  is  of  normal  size,  whilst 
in  Auchmeromyia  it  is  more  than  half  the  length  of  the  whole  abdomen,  and  in  the 
male  the  eyes  are  holoptic  or  close  together,  whilst  in  Auchmeromyia  they  are  wide 
apart.  The  flies  of  this  genus  (three  so  far  described)  attack  man  in  their  larval 
stage  (anyway  two  of  the  three),  and  also  dogs  and  other  animals,  by  burrowing  into 
the  skm  and  producing  painful  boils. 

\C.  anthropophaga,  Griinberg,  is  widely  distributed  in  Africa,  extending  from 
Senegal,  where  its  maggot  is  known  as  the  "ver  de  Cayor,"  and  is  referred 
to  on  p.  590  as  Ochromyia  anthropophaga,  E.  Blanchard,  to  Natal,  where  it  is 
known  as  the  "  Natal  worm,"  and  referred  to  erroneously  on  p.  591  as  Bengalia 
depressa,  Walker. 

[It  is  a  thick-set  Muscid  of  a  general  straw-yellow  colour,  with  blackish  markings 
on  the  dorsum  of  both  thorax  and  abdomen,  about  9*5  mm.  long.  The  larva  is  fat 
and  when  mature  about  12  mm.  long,  bluntly  pointed  in  front,  truncate  behind  ; 
from  the  third  to  eleventh  segments  it  is  thickly  covered  with  minute  recurved 
spines  of  a  brownish  colour,  arranged  in  transverse  series  of  groups  of  two  or  more, 
which  form  more  or  less  distinct  irregular  transverse  rows.  On  each  of  the  two 
posterior  stigmatic  plates,  the  respiratory  slit  on  either  side  of  the  median  one 
is  characteristically  curved,  resembling  an  inverted  note  of  interrogation.  The 
puparium  is  brown  to  ferruginous  or  black  and  about  10  mm.  long.  The  maggots 
are  found  in  both  natives  and  white  men,  and  occur  as  a  severe  pest  in  dogs,  also 
in  monkeys,  rats,  and  other  mammals.  In  Sierra  Leone  it  is  called  the  "  tumba 
fly."  The  larvae  have  been  frequently  found  as  true  subcutaneous  parasites,  each 
larva  living  singly  and  forming  a  boil  or  warble  in  the  skin,  with  an  opening  just  as  in 
an  ox-warble,  through  which  the  maggot  breathes  and  eventually  escapes.  Although 
they  more  usually  occur  as  isolated  specimens,  Marshall  found  in  Salisbury, 
South  Rhodesia,  that  sixty  were  extracted  from  one  lady,  and  Berenger-Feraud,  in 
Senegal,  that  more  than  300  occurred  in  a  single  spaniel  puppy. 

[Neave  {Bull.  Ent.  Res.,  1912,  iii,  p.  217)  records  it  from  ulcers  in  a  native 
at  Lourengo  Marques  in  1908,  and  at  the  same  time  from  ulcers  in  a  dog,  and  that 
it  is  a  severe  pest  to  man  in  Mozambique  and  parts  of  the  Transvaal.  It  seems 
to  be  more  abundant  in  North  Rhodesia  and  Nyasaland  than  to  the  north  (Neave, 
Bull.  Ent.  Res.,  191 2,  iii,  p.  310).  It  is  also  recorded  in  Zanzibar,  German  East 
Africa,  Uganda,  East  Tropical  Africa  (Neave). 

[Simpson  {Bull.  Ent.  Res.,  iii,  p.  170)  records  a  Muscid  larva  taken  from  the 
breast  cf  a  European  in  South  Nigeria  that  was  probably  Cordylobia. 

[It  is  not  known  how  infection  takes  place.  Neave  {Bull.  Ent.  Res.,  iii,  p.  310) 
says  :   "  Many  instances  in  human  beings  would  preclude  the  possibility  of  eggs 


LUND  S    LARVA,   AUCHMEROMYIA    LUTEOLA 


593 


having  been  laid  direct  on  the  skin  :  in  these  cases  they  have  probably  been  laid 
on  the  clothing  put  out  to  dry. 

[Gedoelst  has  described  another  species,  C.  rodhani^  and  Austen  a  third  species, 
C.  prcr^ra7idis^  from  Nyasaland,  Cape  Colony,  Transvaal,  Natal,  North-west 
Rhodesia,  and  German  East  Africa. 

[The  following  are  some  papers  dealing  with  this  subject  :  Proc.  Ent.  Soc, 
London,  for  year  1907,  p.  xlvii  ;  Jomit.  R.A.M.C.,  1908,  pp.  5-1 1,  figs,  i  and  2, 
by  Austen;  Journ.  R.A.M.C.^  1908,  pp.  i  and  2,  by  Major  F.  Smith  ;  Trans.  Soc. 
Trop.  Med.  and  Hyg..,  1910,  iii,  pp.  223-225,  by  Austen. — F.  V.  T.] 


Lund's  Larva. 

Endemic  in  the  region  of  the  Congo  State ;  called  after  Commander  Lund,  from 
the  skin  of  whose  arm  it  was  extracted  ;  12*5  mm.  long,  4*5  mm.  broad  ;  colour 
yellowish,  with  brown  rings,  on  account  of  the  division  of  the  brown  spines  ;  head 
cone-shaped,  with  two  hemispherical  smooth  antennae,  two  thick  black  mouth  hooks 
and  wart-shaped  bodies,  between  which  are  situate  two  to  three  longitudinal  rows 
of  dark  brown  chitinous  laminas.     The  body  segments  are  covered  over  their  whole 


Fig.  411. — Lund's  larva  :  on  the  left,  the  whole  larva,  magnified  six  times.     On  the  right, 
the  head  end,  much  enlarged.     (After  Gedoelst.) 

surface  with  irregularly  distributed  triangular  yellow  spines,  the  points  of  which  are 
coloured  dark  brown.  Its  size  increases  from,  the  second  to  the  sixth  segment, 
diminishes  from  the  seventh  to  the  ninth,  at  the  tenth  it  is  reduced,  and  at  the 
eleventh  quite  small.  The  posterior  stigmata  are  bean-shaped,  each  with  three 
markedly  tortuous  openings.  Duration  of  the  larval  stage  unknown  ;  the  same 
applies  to  the  pupal  and  imago  stages. 

Auchmeromyia  luteola,  Fabricius. 

[This  fly,  the  parent  of  the  so-called  Congo  floor  maggot,^  belongs  to  a  nearly 
allied  Muscid  genus  to  Cordylobia,  but  which  can  at  once  be  told  by  the  great 
length  of  the  second  abdominal  segment.     The  maggot  occurs  in  numbers  in  the 


'  Dutton,  Todd  and  Christy,  "The  Congo  Floor  Maggot,"  Metn.  xiii  Liv.  Sch.   Trop. 
Med.,  p.  40. 


5Q4  THE   ANIMAL   PARASITES   OF   MAN 

native  huts  in  the  Congo  region  and  is  fairly  common  in  central  and  northern  parts 
of  Mozambique  ;  it  is  also  recorded  from  the  Zambesi  River  and  the  vicinity  of 
Barberton  in  the  Eastern  Transvaal  {Bull.  Ent.  Res.,  191 2,  iii,  p.  216),  in  German 
East  Africa,  in  Nyasaland,  and  British  East  Africa.  It  is  also  recorded  from 
Bara,  Kordofan,^  where  they  occurred  on  the  floor  of  the  men's  prison  and  bit  the 
prisoners.  They  were  destroyed  by  sprinkling  Jeyes'  fluid  on  the  floor.  Neave 
states  {ibid.,  p.  310)  that  it  occurs  in  the  more  neglected  huts  in  native  villages 
throughout  tropical  Africa,  and  frequently  enters  a  tent  when  pitched  near  a  village. 
It  is  also  found  in  West  Africa.  The  fly  is  thick-set  and  about  the  size  and  build  of 
a  bluebottle  fly;  length  10  to  12  mm.;  tawny  in  colour  to  dirty  yellowish-brown, 
with  dusky  hairs,  giving  it  a  smoky  appearance  ;  the  flattened  thorax  has  long  dark 
stripes  and  the  abdomen  a  dusky  line  in  the  centre  of  the  second  segment,  which 
meets  a  dark  line  on  its  posterior  border  ;  the  dusky  third  segment  has  a  narrow 
yellowish  anterior  line;  the  fourth  segment  is  also  dusky;  legs  bufl"wiih  black  hairs  ; 
the  fifth  tarsal  segment  black.  The  larviie  are  whitish,  becoming  reddish  after  a  feast 
of  blood,  with  much  wrinkled  skin  and  rather  flat  and  broad.  They  live  in  crevices 
of  the  mud  floor,  under  sleeping  mats  during  the  daytime,  and  come  out  at  night  and 
suck  the  blood  of  sleepers  and  then  retire  to  shelter  again.  Dutton,  Todd,  and 
Christy  noticed  that  where  people  slept  on  beds  or  platforms  raised  above  the  floor 
the  maggots  were  not  so  numerous  as  under  the  sleeping  mats  laid  on  the  ground. 
They  turned  up  many  of  the  maggots  from  a  depth  of  three  inches  or  more.' 
— F.  V.  T.] 

Family.     Oestridae. 

[The  family  of  Oestridcp  or  warble  flies  are  all  parasitic  in  their  larval  stage, 
usually  termed  the  "bot"  stage.  They  are  found  as  parasites  in  warm-blooded 
animals,  and  man  is  frequently  attacked  by  them.  The  members  of  this  family  have 
the  mouth  rudimentary,  many  of  them  are  hairy  and  bee-like,  with  large  eyes  and  the 
head  large,  the  lower  part  more  or  less  swollen.  The  thorax  is  large  with  a  distinct 
transverse  suture,  and  the  abdomen  short  and  stumpy  or  very  slightly  elongated. 
The  male  genitalia  are  hidden,  whilst  the  female  ovipositor  is  often  elongated.  The 
wings  may  be  transparent  (Hypoderma)  or  mottled  (Gastrophilus),  and  have  muscid- 
like  venation  ;  the  tegulas  usually  large,  the  legs  moderately  long. 

[As  a  rule  each  species  is  confined  to  a  particular  host,  but  as  we  see  recorded 
here  those  that  attack  animals  may  also  attack  man.  The  flies  occur  in  warm  weather 
and  usually  during  the  warmest  part  of  the  day,  and  have  a  strong  dislike  to  shade 
and  water.  The  genus  Hypoderma  attack  oxen,  sheep,  goats,  antelope  and  musk 
deer;  Oestrus,  sheep,  antelope  and  horses;  Gastrophilus,  the  horse  and  ass; 
Cephenomyia,  the  deer  ;  Cepholomyia,  the  camel  and  bufifalo  ;  Dermatobia,  dogs, 
cats,  oxen,  deer,  apes  and  man  ;  Cuterebra  and  Rogenhofera,  rodents  and  opossums- 

[Some  live  as  parasites  in  the  stomach  and  intestines  (Gastrophilus)  ;  others 
infest  the  skin  (Hypoderma,  Dermatobia  and  Oestromyia,  the  latter  on  Lagomys  and 
Hypodaus) ;  (Edemagena  tarandi  also  infests  the  skin  of  the  reindeer  in  Siberia  and 
boreal  America.  Oestrus  lives  in  the  nasal  sinus,  and  Cephalomyia  in  the  throat  as 
well,  Cuterebra  and  Rogenhofera,  the  skin  or  scrotum,  so  that  we  have  really  three 
groups  of  parasitic  oestride  larvae  :  (i)  cutaneous,  (ii)  intestinal,  and  (iii)  facial. 

[No  species  seems  confined  to  man,  but  the  so-called  "creeping  disease,"  caused 
by  Hypodermae,  and  the  attack  of  sheep  nasal  fly  are  comparatively  common,  as  also 
is  the  Dermatobia  attack.— F.  V.  T.] . 


'  Balfour,  Journ.  Trop.  Med.,  1909,  xii,  No.  4,  p.  47. 
2  Journ.  Trop.  Med.,  1905,  viii,  No.  6,  p.  90. 


cutaneous  oestrid^:  595 

Cutaneous  Oestrid^. 

The  eggs  are  deposited  on  the  surface  of  the  body  ;  the  larvns  burrow  in  the 
skin,  which  they  reach  after  somewhat  long  peregrination. 

Genus.     Hypoderma,  Latreille, 

Hypoderma  bovis,  de  Geer. 

The  cattle  fly  or  warble  fly,  which  swarms  during  the  hot  season,  settles  on  the 
head  or  on  the  hair  of  grazing  cattle  ;  through  the  young  being  licked  off  they  gain 
access  to  the  mouth  and  are  swallowed.^  The  larvae  appear  first  in  the  commencing 
portion  of  the  stomach,  to  escape,  as  some  state,  into  the  preceding  sections  of  the 
alimentary  canal ;  at  any  rate,  they  are  found  from  July  onward  regularly  in  the 
submucous  tissue  of  the  pharynx,  in  which  they  travel  about  for  several  months 
(up  to  November,  and  in  isolated  cases  up  to  February)  ;  they  then  penetrate  the 
muscularis  and  migrate  by  way  of  the  subserosa  along  the  mediastinum,  the  crura 
of  the  diaphragm,  the  renal  capsules,  and  the  intermuscular  connective  tissue  of  the 
psoas  muscle  in  the  direction  of  the  spinal  canal,  into  which  they  penetrate  by  way 
of  the  muscles  and  nerves,  through  the  intervertebral  foramina.  Here  they  stay  for 
about  two  to  three  months,  then  they  leave  the  spinal  canal  again  through  the 
vertebral  foramina  and  make  their  way  (from  January  to  March)  through  the  inter- 
muscular connective  tissue  of  the  muscles  of  the  back  to  the  skin  of  the  back,  where 
sooner  or  later  (from  January  to  June)  they  arrive  and  enter  a  resting  stage,  which 
commences  with  penetration  of  the  skin  and  terminates  with  outward  migration 
from  the  boils  due  to  the  wound  set  up  by  the  maggot.  At  the  commencement 
of  this  period  the  larvae  cast  their  skin,  and  their  form,  hitherto  cylindrical,  becomes 
oval.  After  about  a  month,  a  second  moulting  of  the  skin  takes  place — the  third 
larval  stage,  which  lasts  about  two  and  a  half  months  (up  to  June).  The  approaching 
end  of  the  same  is  indicated  by  a  change  of  colour  on  the  part  of  the  larva  from  the 
hitherto  yellowish-white  to  brown  and  finally  to  blackish-brown.  When  they  have 
become  mature  the  larvre  leave  the  warbles,  drop  on  to  the  ground  and  pass  into 
the  pupal  stage  in  the  superficial  layers  of  the  soil  within  twelve  to  thirty-six  hours. 
After  about  a  month  the  flies  emerge.  Irregularities  with  regard  to  the  time  and 
direction  of  the  migrations  of  the  larvss  take  place  (Jost,  H.,  in  Zeitschr.  f.  wiss. 
ZooL,  1907,  xxxvi,  p.  644). 

In  a  number  of  cases  the  larva  of  the  cattle  fly  has  been  observed 
in  the  human  integument,  usually  in  the  winter  months,  that  is, 
during  the  migration  period  ;  consequently,  it  is  not  surprising  that 
the  larvae  before  they  enter  on  the  resting  stage  and  produce  a  warble 
undergo  migrations.  But  that  this  takes  place  subcutaneously — 
w^hich  does  not  appear  to  be  so  in  the  case  of  cattle — is  perhaps 
explained  by  the  fact  that  in  man,  on  account  of  the  short  space  that 
has  to  be  traversed,  the  larvae  are  not  sufficiently  developed  to  enter 
on  the  resting  stage  simultaneously  upon  having  obtained  access 
to  the  integument.  Whether  the  Oestrid  larvae  in  Bulgaria  that 
similarly  migrate  beneath  the  skin  in  man  belong  to  the  cattle  fly 
or   to    another   species,  or   even    another   genus,  has   not   yet   been 

1  [This  is  not  the  case,  for  Carpenter  has  shown  that  muzzled  calves  become  infected 
("Mem.  First  Int.  Cong.  Ent.,"  pp.  289-293).  ]o%\.{Zeiischr.  f.  wiss.  ZooL,  1907,  xxxvi, 
pp.  644-715)  thinks  that  the  ova,  not  young  larva;,  are  ingested  (vide  note  in  Supplement. 
— F.  V.  T.J- 


596  THE    ANIMAL   PARASITES   OF   MAN 

ascertained.  (Doctorow,  in  Arch,  de  Par.,  1906,  x,  p.  309  ;  Spring,  A., 
in  Bull.  Acad.  set.  Belg.,  1861  (2),  iv,  p.  172;  Walker,  R.,  in  Brit.  Med. 
yoiirn.,  1870,  i,  p.  151  ;  Kjelgaard,  in  IJgeskr.  f.  Laeger,  1904,  p.  535  ; 
Condorelli,  M.,  in  Bull.  Soc.  Zool.  Italy  1904,  xiii,  p.  171.) 

Hypoderma  lineata,  de  Villers. 

The  larvie  of  this  species,  that  occurs  not  only  in  Europe  but  in  North  America, 
live  under  similar  conditions  in  the  skin,  very  rarely  in  man;  also  migrating 
subcutaneously  (Topsent  in  Arch,  de  Par.,  1901,  iv,  p.  609). 

[In  Sweden,  the  ox  warble  fly  {H.  bovis)  is  well  known  to  attack  man. 
Schoyen  states  "  that  over  100  years  ago  up  to  the  present  time  cases  of  travelling 
grubs  under  the  human  skin  in  some  districts  of  Sweden  were  well  known." 
The  species  appeared  to  be  H.  bovis,  many  of  which  he  had  examined.  They 
accomplished  long  ramblings  under  the  skin,  always  in  an  upward  direction, 
previous  to  their  appearance  through  an  opening  in  a  tumour  on  the  upper  part  of 
the  body,  on  the  head,  neck,  or  shoulders.  An  interesting  case  is  recorded  in 
Insect  Life.,  ii,  pp.  238-239.  A  bot  similar  to  H.  diafia  was  taken  from  the  eye 
and  cheek  of  a  child  at  Kane,  McKean  County,  Pa.,  U.S.A.  It  was  said 
to  have  travelled  in  five  months  from  the  elbow  to  the  eye.  Riley  later  {Insect 
Life,  iv,  p.  31c)  was  inclined  to  think  the  maggot  was  that  of  H.  lineata, 
the  common  American  ox  warble,  which  is  also  found  in  Europe  in  great 
numbers.  I  have  recorded  another  case  in  England  {Rept.  Econ.  Zool.  for 
year  ending  September  30,  1910,  p.  128),  where  Dr.  Menzies  removed  the  larva 
of  H.  bovis  from  the  upper  eyelid  of  a  patient.  It  caused  considerable  swelling 
of  the  face,  much  pain  and  distress  ;  but  the  case  did  well,  and  the  wound  healed 
at  once.  The  larva  was  nearly  mature.  Numerous  other  references  to  this 
so-called  creeping  disease  will  be  found  in  the  Supplement. 

[It  is  quite  probable  that  bovis  and  lineata  are  confused  in  the  latter  accounts. 
The  larvae  are,  however,  easily  distinguished  if  carefully  examined. — F.  V.  T.] 

Hypoderma  diana,  Brauer. 

In  its  larval  stage  it  lives  like  other  species  of  Hypoderma,  attacking  the  red 
deer  {Cervas  elaphas)  and  roe  deer  {Cervas  capreolus)  ;  it  is  occasionally  also  found 
in  man  (Joseph,  in  "Myiasis  externa  dermatosa,"  Hamburg,  1800;  Volkel,  in 
Berl.  klin.    IVochenschr.,   1883,  ^x,  p.   209). 

Genus.     Dermatobia,  Brauer. 
Dernnatobia  cyan  iventris,  Macq. 

Syn. :  Dermatcbia  noxialis,  J.  Goudot. 
The  genus  Dermatobia  represents  the  subcutaneous  Oestridce  of  Europe  in 
warmer  parts  of  America.  Both  domesticated  and  wild  mammals  are  attacked, 
according  to  one  statement  birds  also  {Ramphastus),  and  man  with  fair  frequency.^ 
It  is  assumed  that  in  all  cases  one  and  the  same  species  is  concerned,  for  which 
recently  a  name  originating  from  C.  Linne,  jun.  {Oestrus  ho?ninis),  has  been 
employed.     Three  larval  stages  are  recognized  in  the  skin  ;  the  two  first  appear  to 

Duprey  advances  the  opinion  that  Dermatobia  deposits  its  eggs  not  only  on  the  skin  of  man 
and  animals,  but  also  on  the  leaves  and  twigs  in  the  bush,  where,  too,  young  larvie  have  been 
met  with  which  gain  access  from  hence  to  men  and  animals  {Jonrn.  Trop.  Med.  and  Hyg.,  1906). 


DERMATOBIA   CYAXIVENTRIS 


597 


resemble  one  another  in  the  club-shaped  or  tadpole-like  appearance  (called  macaque 
in  Cayenne,  mayacuil  [mayoquil]  in  Mexico),  the  third  is  swollen  spindle-shaped 
(Berne,  called  torcel).  Segments  2  to  4  in  the  club-shaped  larvae  are  closely  beset 
with  small  black  spines,  segments  5  to  7  bear  at  the  anterior  border  a  complete  ring 
of  strong  black  hooks,  segments  4  to  6  a  similar  ring,  which,  however,  is  interrupted  at 
the  ventral  surface.  The  four  last  segments  forming  the  tail  are  smooth,  only  at  the 
posterior  end  are  there  small  spines.     The  arrangement  of  spines  of  the  third  stage 


Fig.  412. — Dervialobia  noxialis,  Goudot. 


Fig.  413. — Larva  of  Der- 
viatobia  cyaniventris  in  its 
natural  size  and  magnified. 
(After  Blanchard.) 


Fig.  414.  —  Larva  of  Der- 
matohia  cyaniventris.  Enlarged' 
(After  Blanchard.) 


differs  from  this.  Italian  workmen  that  have  been  employed  in  Brazil  show  the 
presence  of  Dermatobia  larvae  on  their  return  (Blanchard,  in  Bull.  Soc.  Ent. 
France,  1893,  P-  24;  Bull.  Soc.  centr.  de  Med.  vet.,  1896;  Ann.  Soc.  Ent. 
Fra?tce,  1894,  Ixiii,  p.  142  ;  Ward,  H.  B.,  in  Mark  Annivers.  Vol.,  Article  25, 
p.  483,  New  York,   1903). 

l^Dennatobia  cya7iiventris,  Macquart,  1843,  is  said  not  to  be  the  same  as  noxialis 
{vide  Brauer,  "  Mono.  Oestriden,"  1863,  p.  266).     It  is  known  by  various  other  names, 


598  THE    ANIMAL    PARASITES    OF    JMAN 

as  niiche  or  giisano  in  New  Granada,  the  ura  in  Brazil,  and  the  macaw  fly  in 
Cayenne.  It  occurs  in  Central  and  South  America  and  the  West  Indies.  According 
to  Goudot  the  fly  is  found  in  great  numbers  on  the  borders  of  large  woods  and 
lands  covered  with  underwood. 

[It  is  seldom  that  more  than  one  larva  is  found  in  each  individual.  It  is  generally 
found  in  the  arm  and  leg,  but  now  and  then  the  face.  The  perfect  insect  has  never 
been  bred  from  a  larva  removed  from  a  human  being,  so  that  there  is  still  uncertainty 
as  to  the  actual  species.  D.  cyanive?itris  is  11  to  12  mm.  long,  has  an  ochraceous 
buff-coloured  face,  dark  grey  thorax,  metallic  dark  blue  to  purple  abdomen,  and 
brownish  wings.     D.  noxialis  is  somewhat  larger. 

[In  the  Journal  of  Tropical  Medicine  and  Hygietie,  January  15,  1905,  viii,  p.  23, 
reference  is  made  to  this  Oestrid  in  Trinidad,  where  it  is  called  the  "mosquito 
worm."  One  case  here  recorded  showed  no  fewer  than  four  worms  on  the  chin  and 
one  on  the  hand.  It  is  here  stated  that  the  fly  never  attacks  man  or  animals  directly, 
as  it  is  said  to  do  by  Scheube,  but  that  the  eggs  are  deposited  on  leaves  and  branches 
in  wooded  lands  and  forests,  and  thus  man,  hunting  dogs  and  wild  animals  in 
passing  through  get  the  larvae  deposited  on  them  accidentally.  The  affection  is 
common  in  Trinidad.  Mention  is  made  that  a  little  i  in  40  carbolic  lotion  syringed 
into  the  aperture  in  the  skin  over  the  worm  quickly  killed  it. 

[The  cattle  worm,  or  founzaia  ngombe,  is  the  name  given  to  a  larva  which 
develops  beneath  the  skin  of  oxen  and  men  in  Central  Africa,  especially  amongst  the 
natives  and  stock  of  Unyamonezi.  According  to  P.  Dutrieux,  the  egg  is  laid  by  a 
large  fly  that  accompanies  cattle.  It  is  unknown  between  the  central  plateau  or  the 
Ugogo  and  the  East  Coast.— F.  V.  T.] 

Cavicolous  Oestrid^. 
The  forms  belonging  to  this  group  inhabit  as  larvae  the  nasal  and 
frontal  sinuses  of  ruminants,  Eqnidce  and  Prohoscidcc,  which  they  leave 
for  the  pupal  stage.     The  larva  of — 

Genus.     Oestrus,  Linnaeus, 

Oestrus   (Cephalomyia)    ovis,  L., 

occurring  in  sheep,  has  also  been  observed  in  man  in  six  cases  in  the 
nose  and  larynx  (Saitta  in  Gaz.  d.  Osp.  d.  Clinic,  1903,  No.  128).  So 
far  as  is  known,  the  eggs  are  deposited  in  the  nasal  cavity. 

{^Oestrus  ovis  frequently  occurs  in  man.  MM.  Sergent  (^«;/.  de  VInst.  Pasteur, 
1907,  pp.  392-399)  mention  that  they  lay  their  ova  on  the  noses,  eyes  and  mouth  of 
humans  in  Algeria  whilst  flying,  but  that  they  disappear  after  three  to  ten  days  or 
the  inflammation  produced  by  them.  Portschinsky  {Mem.  Bur.Ent.  Sci.  Com.  Cent. 
Bd.  Land  Adm.  and  Agric,  1913,  x.  No.  3,  p.  63)  also  gives  cases.  He  doubts 
that  ova  are  laid  on  the  nose  ;  evidently  the  Russian  habit  is  anomalous,  for  the 
Sergents,  Collings  and  myself  find  ova  laid  as  a  common  occurrence.  I  have  often 
seen  them  on  the  nose  of  sheep.  This  fly  also  occurs  in  the  Argentine  (Serres,  in 
Gaceta  Rural,  April,  1913,  vi,  pp.  759-761). 

[The  tamne  or  thimni  of  the  Kabyles,  a  human  myiasis  of  the  Tuareg  mountains 
in  the  Sahara,  is  caused  by  Oestrus  ovis.  Here  the  larvcE  are  said  to  be  ejected  on 
to  the  conjunctival  and  nasal  mucous  membrane  of  humans. 

[Ed.  and  Lt.  Sergent  [Bull.  Soc.  Path,  e.xot.,  191 3,  vi,  No.  7,  pp.  487-488)  report 
their  attack  from  the  Ahaggar  mountains,  in  Central  Sahara.  The  Tuareg  name  for 
.the  fly,  tamne,  is  the  Targui  form  of  the  word  thimni  used  by  the  Kabyles.— F.  V.  T.] 


GASTRICOLOUS   OESTRID^ 


599 


GASTRICOLOUS   OESTRID^. 

The  eggs  are  deposited  on  the  hairs  of  Eqiiidcv,  and  the  larvae 
escaping  from  them  are  Hcked  up  and  swallowed.  They  pass  their 
larval  stage,  according  to  the  species,  in  various  parts  of  the 
intestine  and  stomach,  and  when  mature,  pass  out  per  aiiiim  in 
order  to  undergo  the  pupal  stage. 

Genus.     Gastrophilus,   Leach. 

One  of  the  most  frequent  species  is  Gastrophilus  eqiii^  Fabr. ;  the  eggs  are  laid 
on  the  hairs  ;  the  larvae  live  some  ten  months  in  the  stomach,  living  attached 
to  the  inner  surface.  The  eggs  of  G.  hceniorrhoidalis,  L.,  are  deposited  on 
the  lips  or  the  long  hairs  on  them.  The  larvae  adhere  to  the  cardiac  end  of 
the  stomach,  to  the  stomach  itself,  and  finally  to  the  terminal  portion  of  the 
intestine.  Here,  however,  and  elsewhere  in  the  intestine,  the  larva:  of  G.  pecoruni', 
Fabr.,  are  also  niet  with,  whilst  the  larva:  of  G.  nasalis  (so  called  because  the 
eggs  are  deposited  in  the  nasal  orifices)  almost  exclusively  inhabit  the  anterior 
section  of  the  duodenum. 

Cholodkowsky  attributes  the  "  wormlet "  observed  by  Samson  and  Sokolew 
{Wratsch,  1895,  Nos.  48  and  57)  and  others  {ibid.,  1896-98)  to  Gastrophilus  larvae. 
It  burrows  into  the  epidermis  of  man  by  minute  passages.  This  observation 
should,  however,  be  verified.  The  phenomenon  is  designated  as  skin- mole,  larva 
migrans,  and  creeping  eruption. 

Other  Papers  on  Dipterous  Larv/e,  etc.,  in  Man. 
(i)  "  Ein   Fall   von    lebenden    Fliegenlarven   im    menschlichen    Magen,"   Deutsch   nied. 
Wochenschr.,   Leipz.    and  Berl.,  xxiv  (12),  pp.   193-194.       Bachmann,  and   review  of  same, 
"  Living  Fly  Larvre  in  the  Human  Stomach,"  fhiladelphia  Med.  fota-n.,  1898,  i,  18,  p.  773. 

(2)  "Siidi  una  larva  di  dittero  parassita  della  congiuntiva  umana,"  Ami.  di  ottal.,  Paira, 
1895,  xxiv  (4),  pp.  329-336,  I  fig.,  E.  Baquis. 

(3)  "  Sur  quelques  dipteres  suceurs  de  sang,  observe  a  Terre-Neuve,"  Arch,  de  Par., 
Patis,  1900,  iii  (i),  pp.  202-204,  E.  Barret. 

(4)  "An  Account  of  the  Larvae  of  two  Species  of  Insects  discharged  from  the  Human 
Body,"  Edin.  Med.  and  Surg,  /ourn.,  January  I,  i8n,  vii  {25),  pp.  41-48,  I  pi.,  figs.  I  to  8, 
T.  Bateman. 

(5)  "  Un  cas  de  myiase  par  la  Sarcophaga  magnifica  en  Roumanie,"  Bull.  Soc.  ZooU 
de  France,  Par.,  189 1,  xvi  (2),  pp.  25-26,  R.  Blanchard. 

(6)  "  Sur  les  oestrides  americains  dont  la  larve  vit  dans  la  peau  de  I'homme,"  Ann.  Soc. 
ent.  de  France,  1892,  v,  pp.  109-154,  figs.  It  12,  R.  Blanchard. 

(7)  "  Note  additionnelle  sur  les  oestrides  americans  dont  la  larve  vit  dans  la  peau  de 
I'homme,"  Bull.  Soc.  ent.  de  France,  Pari--,  1894,  xiv,  pp.  209-211,  R.   Blanchard. 

(8)  "  Note  sur  des  larves  de  Dermatobia  provenant  de  Bresil,"  Bull.  Soc.  ent.  de  France, 
Paris,  1893  {2),  pp.  24-27,    R.  Blanchard. 

(9)  "  Larven  der  Wohlfahrtfliege  {Sarcophila  wolfahrtii)  im  Zahnfleische  eines  Menchen," 
Wratsch.,  St.  Petersburg,  1888,  5-6,  E.  K.  Brandt. 

(10)  '•  Ueber  den  sogenannten  Oestrus  honiinis  und  die  oftmals  besichteten  Verirrungen  von 
Oestriden  der  Saugetheieie  zum  Menchen,"  Verhandl.  d.  k.  zool.-bot.  Gesellsch.,  i860,  x 
Abhandl.,  pp.  57-72,  Brauer. 

(11)  "Ueber  die  Larven  der  Gattung  Cuterebra,  Clk.,"  Verhandl.  d.  k.  zool.-bot. 
Gesellsch.,  i860,  x  Abliand.,  pp.  777-786,  Brauer. 

(12)  "  Des  desordres  produits  chez  I'homme  par  les  larves  de  la  Lucilia  hominivorax,''^ 
These,  Paris,  1864,  43  pp.,  V.  Audouit. 

(13)  "  Note  on  the  'Flesh  Worm,'"  Med.  Press  and  Circ,  London,  April  12,  1882, 
Ixxxii  (N.S.  xxxiii),  p.  314,  P.  S.  Abraham. 


OOO  THE   ANIMAL   PARASITES    OF   MAX 

(14)  *'  Larvas  de  la  Calliphora  liniensis  en  fosas  nasalis,"  1855,  18  pp.,  F.  Aguirre. 

(15)  "  Raro  caso  di  parasilismo  nell  'uomo  dovuto  alia  larva  di  una  mosca  {Sarcophaga 
affinis,  Meigen),"  Boll.  d.  Soc.  Rom.  fer  gli  Stud.  Zool.,  Roma,  1893,  i^  (5-6),  pp.   278-289, 

I  pi.,  3  figs.,  Giulo  Alessandrini. 

(16)  "  Observations  sur  I'espece  de  ver  nomme  Macaque  (Oestrus),"  Mem.  Acad.  Sci- 
par  Hist.,  1753,  p.  72,  F.  Artur, 

(17)  "  Contribuicao  ao  estudo  da  biologia  da  Dermatobia  cyanivent?-is,'^  Trav.  do  Inst,  de 
Maiigufjihos,  1908. 

BlTlNG-MOUTHED    AND   OTHER    NOXIOUS    DlPTERA   WHICH    MAY   BE 

Disease  Carriers. 

[Amongst  the  division  Brachycera  (as  meant  in  this  work)  we  get  several  groups 
of  flies  which,  like  the  fleas  and  mosquitoes,  are  partially  parasitic  on  man,  the  adults, 
mainly  in  the  female  sex,  being  provided  with  a  piercing  mouth  with  which  they 
extract  the  blood  of  man  and  animals.  The  importance  of  these  parasites  is  net  the 
mere  fact  that  they  feed  upon  our  blood,  but  that  they  often  carry  germs  from  man 
to  man  (tsetse-flies  and  trypanosomiasis,  Tabaiiidce  and  anthrax).  Amongst  the 
most  important  biting-mouthed  Diptera  in  this  section  are  the  following  :  Tabanidcp., 
or  gad-flies  ;  Glossitice,  or  tsetse-flies  ;  and  certain  other  Muscidcr.  Some  of  the 
exotic  Asilidic  and  a  few  Leptidcr  also  bite  man. 


Family.     Tabanidae   (Gad-flies). 

[The  Tabanidce  have  a  broad,  rather  flattened  body  and  a  large  head  ;  eyes  united 
in  the  male  (except  in  some  Chrysops).  The  antennae  are  composed  of  three  seg- 
ments, have  the  third  joint  composed  of  five  to  eight  annuli — in  Chrysops  they  are 
fairly  long.  The  proboscis  is  projecting,  and  sometimes  much  elongated.  The  legs 
are  moderately  stout.     The  venation  of  the  wings  is  shown  in  fig.  415. 

[This  family  of  gad  or  horse  flies  contains  a  great  number  of  genera,  all  of 
which  may  bite  animals  and  man  more  or  less  severely.  The  female  alone  is 
blood-sucking,  the  males  feed  upon  the  juices  of  flowers.  The  females  deposit  their 
spindle-shaped  white,  black,  or  brown  eggs  on  leaves,  stems  of  plants  that  either 
overhang  or  stand  in  water,  and  amongst  rushes  ;  they  are  at  first  white,  but  become 
brown  or  black.  The  eggs  are  laid  in  rounded,  flattened  or  conical  masses  com- 
posed of  layers  one  upon  the  other.  The  larvne  are  carnivorous,  feeding  upon  snails, 
worms,  other  larvce,  etc.,  and  have  a  distinct  head ;  they  are  cylindrical,  com- 
posed of  eleven  segments,  the  last  with  a  vertical  breathing  pore,  or  the  last  two 
segments  may  form  a  breathing  tube.  The  majority  taper  to  a  point  at  each  end,  in 
colour  shining  white  or  dull  grey  to  yellowish,  many  of  the  larger  specimens  mottled 
or  banded  with  dark  brown  or  black.  The  first  seven  abdominal  segments  are 
encircled  near  the  anterior  margin  with  a  ring  of  fleshy  protuberances  consisting  of 
a  transverse  dorsal  ridge  which  may  be  divided  by  a  depression  into  two.  The 
young  larvae  burrow  into  any  soft  vegetable  substance  ;  they  live  both  in  the  water 
and  under  damp  soil  surrounding  water,  also  in  damp  earth  generally.  The  larvte 
are  not  only  carnivorous,  but  they  are  cannibals,  frequently  devouring  their  own 
species.     They  may  take  more  than  a  year  to  mature. 

[The  pupcx  are  found  close  to  the  surface  of  mud  and  earth,  and  are  mostly  dull 
yellowish  to  brown  in  colour,  with  rows  of  spines  on  the  distal  third  of  each 
abdominal  segment ;  the  thorax  bears  a  pair  of  ear-shaped  spiracular  structures,  and 
there  are  also  six  denticles  at  the  apex  of  the  abdomen. 

[A  habit  common  to  the  adults  of  most  of  the  Tabaiiidce  of  considerable  economic 


TABANIDi^i: 


6oi 


importance  is  that  of  the  adults  coming  to  water  to  drink.  Portschinsky'  has  found 
that  by  applying  kerosene  to  the  pool  they  frequent  the  adults  are  killed,  and 
Hine-  that  the  same  oil  kills  the  larvas  that  fall  into  the  water  from  eggs  laid  on 
plants  above, 

\Tabanida'  are  not  only  of  importance  as  purely  biting  insects,  for  they  may 
often  convey  pathogenic  organisms  from  one  animal  to  another,  such  as  the 
bacillus  of  anthrax,  which  they  are  known  to  carry,  and  possibly  also  trypano- 
somes  in  regard  to  man.  Chrysops  also  acts  as  a  host  of  Filaria  loa  in  South 
Nigeria  (Lei per,  Brit.  Med.  Joiirn.^  January,  191 2,  pp.  39-40).  Two  species  are 
incriminated,  viz.^  C.  silacea  and  C.  dimidiata.  With  animals  these  flies  play 
a  more  important  part,  for  MM.  Sergent,  in  Algeria,  have  proved  that  species  of 
Tabanus  are  able  to  transmit  three  forms  of  animal  trypanosomes  by  biting  a 
healthy  animal    as    long    as    twenty-two    hours    after   having   bitten   an   unhealthy 


Fig.  415. — The  ox  gad  fly  {Tabanus  boviims,  Linn.). 

one.  In  India  they  have  also  been  shown  to  transmit  the  parasite  of  "surra" 
in  dogs  and  rabbits  by  Rogers.  Other  observers  have  since  corroborated  these 
results,  and  Mitzmain,  who  has  recently  performed  valuable  work  in  this  connection, 
states  that  T.  striaius  is  undoubtedly  the  carrier  of  this  disease  in  the  Philippine 
Islands.  Certain  members  of  the  genus  Haematopota  have  also  been  shown  to  be 
capable  of  the  direct  transmission  of  Trypanosoma  evansi.  Martoglio  {Ann. 
dig.  sper.^  191 3»  xxiii,  N.S.,  No.  3,  pp.  363-366)  states  that  the  trypanosome  disease 
of  dromedaries  known  as  salaf  is  transmitted  by  Tabanidce.,  especially  Pangonia 
{P.  magretti  and  P.  beckeri)  in  Italian  Somaliland.  It  is  quite  likely  that  these 
flies  play  a  much  greater  part  in  the  spread  of  such  diseases  than  is  imagined 
at  the  present  time. 

[The  TabanidcE  are  divided  into  two  groups  or  subfamilies  :  (i)  The  Pangonince.^ 


'  Vide  Btdl.  20,  N.  Sc,  U.S.  Div.  Ent. 

-  '*  Tabamda  of  Ohio,"  Ohio  State  University  Bidt.  19,  1903,  sec.  7,  p.  14. 


6o2 


THE    ANIMAL    PARASITES    OF   MAN 


and  (2)  the  Tabaniiice ;  the  former  have  spurs  on  the  hind  tibia"  and  usually  ocelli  ; 
the  latter  have  neither  tibial  spurs  nor  ocelli. 

[The  Paiigoni7ice  contain  two  main  genera,  Pangonia  and  Chrysops.  In  the 
former  the  proboscis  is  much  elongated,  and  the  third  antennal  segment  is  composed 
of  eight  rings,  and  is  never  angulated  or  ungulated  at  the  base.  The  proboscis 
is  often  very  long. 

[In  Chrysops,  the  so-called  blinding  storm  flies,  all  the  three  segments  of  the 
antennae  are  long,  the  third  having  only  five  annulations,  and  the  proboscis  short 
but  very  strong. 

[There  are  many  genera  in  the  Taba7iince,  which  are  found  in  all  parts  of  the 
world,  of  which  two  only  are  shown  here — viz.^  Tabanus  and  Ha^matopota.  The 
former  has  the  first  two  segments  of  the  antennas  short,  the  third  angulated 
at  the  base,  sometimes  spurred  and  composed  of  five  annulations ;  the  second 
has  the  second  segment  short,  and  the  third  composed  of  four  annulations — 
never  angulated  nor  spurred  at  the  base — and  the  wings  are  adorned  with  grey  or 
brown  markings.  These  latter  are  usually  called  "brimps"  and  "clegs"  in  Britain, 
the  former  gad  or  horse  flies,  the  seruts  and  mangrove  flies  of  tropical  countries. 


Fig.  416.— The  brimp  {HcBinaiopotaphivialis,  Linn.). 


Family.     Asilidae  (Wolf  Flies). 

[These  flies  are  of  little  importance  in  regard  to  the  subject  dealt  with  in  this 
book  ;  but  I  have  notes  sent  concerning  the  biting  habits  of  one  or  more  species 
belonging  to  this  family  from  the  Malay  States  and  Africa. 

{^Asilidce,  or  wolf  tiies,  are  easily  told  by  the  following  characters  :  Large  or 
moderate-sized  flies,  thickly  hairy  ;  head  separated  from  thorax  by  a  narrow  neck  ; 
eyes  separated  in  both  sexes;  proboscis  firm  and  horny,  adapted  for  piercing; 
abdomen  long,  pointed,  and  composed  of  eight  segments.  Legs  strong  and  bristly, 
of  moderate  length.  Wings  sometimes  mottled,  lying  parallel  over  the  abdomen 
when  at  rest.  There  are  nearly  3,000  species.  They  live  mostly  upon  insects,  but 
some  are  said  to  bite  animals  and  man.  They  are,  however,  of  little  importance  in 
this  respect. 


BLOOD-SUCKING   MUSCID^  603 

Family.     Leptidae. 

[This  widely  distributed  family  of  flies  has  a  few  species  which  suck  the  blood  of 
man,  and  the  writer  has  been  personally  badly  bitten  in  Norway  by  a  Leptis  which 
was  apparently  Leptis  scolopacea. 

[The  Leptidce  have  usually  blotched  wings  and  similar  venation  to  Tabanus  ;  they 
are  elongated  flies  of  moderate  or  large  size,  and  of  dull  colours.  The  antennas  are 
varied  and  consist  of  three  segments,  either  with  or  without  a  terminal  bristle  or  with 
the  third  segment  compound,  and  in  a  few  they  may  be  almost  nematocerous.  The 
wing  veins  are  distmct,  very  crowded  anteriorly,  the  third  long  vein  is  furcate,  basal 
cells  large,  and  there  are  usually  five  posterior  cells,  the  anal  cell  being  open  in  some  ; 
the  squanicC  are  always  small,  sometimes  only  rudimentary, 

[Four  are  known  to  be  blood-suckers,  namely  the  American  Symphoromyia, 
Trichopalpus  obscurics  in  Chili,  and  Leptis  strigosa  and  L.  scolopacea  in  Europe. 
The  genus  Symphoromyia  has  a  single  spur  on  the  hind  tibicE,  none  on  the  fore  or, 
raid  tibic\2,  the  third  segment  of  the  three-ringed  antennae  kidney-shaped,  and  a  short 
proboscis.  In  the  genus  Leptis  the  hind  tibiae  have  two  spurs,  and  the  third  antennal 
segment  is  not  reniform. 

[The  other  biting  genus  Trichopalpus  can  be  told  at  once  by  the  elongated 
proboscis.  Most  of  this  family  live  upon  other  insects.  The  larvas  live  in  earth, 
decaying  wood,  sand,  stagnant  waters,  and  the  nests  of  wood-boring  beetles  ;  they 
are  usually  cylindrical  and  may  have  fleshy  abdominal  legs  ;  the  anal  segment  has 
a  transverse  cleft,  and  often  two  posteriorly  directed  processes  and  two  stigmata 
between  them.  They  are  all  predaceous,  and  in  one  genus  (Vermileo)  make  pitfalls 
in  sand  like  the  ant  lions  {Mynneleon). 


Bloods  sucking  Muscidse. 

[The  blood-sucking  MuscidcE  are  mainly  contained  in  the  following  genera : 
Glossina,  Stomoxys,  Haematobia,  Lyperosia,  Stygeromyia,  Philsematomyia  and 
Bdellolarynx. 

[The  first  is  the  most  important  genus  on  account  of  the  part  it  plays  in  the 
spread  of  trypanosome  diseases.  Stomoxys  may  also  serve  as  a  disease  carrier. 
The  remainder  and  a  few  more  genera  cause  considerable  annoyance  by  their 
bites,  and  may  also  act  as  occasional  carriers  of  pathogenic  organisms.  All  these 
flies  have  their  mouth  parts  elongated  to  some  extent,  forming  a  distinct  proboscis, 
which  becomes  more  or  less  strongly  chitinized  ;  the  labella  are  usually  serrated 
or  spiny,  and  thus  form  a  structure  easily  capable  of  piercing  the  skin.  Unlike 
the  Culicidcc^  the  blood-sucking  Muscidce  have  the  sanguinary  habit  common  to  both 
sexes. 

Genus.     Glossina,  Westwood. 

[This  genus  contains  sixteen  species,^  all  of  which  are  confined  to  the  Ethiopian 
region.  Glossina  may  be  distinguished  from  other  allied  genera  by  the  proboscis, 
the  antennae,  wings,  and  male  genitalia.  The  proboscis  projects  forwards  and  has 
1  swollen  bulb-like  base  to  the  slender  labium  which  holds  the  two  structures,  the 
needle-like  epipharynx  and   the  thread-like  hypopharynx  ;   the  whole  proboscis  is 

'  This  does  not  include  G.  mactdata,  Newstead,  which  is  regarded  by  Austen  as  a 
synonym  of  G.  palpalis,  Rob.  Des.  ;  according  to  this  authority  the  curiously  spotted 
appearance  of  the  type  and  only  example  of  G.  maculata  is  due  to  foreign  matter. 

38 


604. 


THE   ANIMAL   PARASITES   OF   MAN 


ensheathed  in  the  maxillary  palpi.  The  antennas  have  the  first  two  segments  small, 
the  third  large  with  a  marked  pore,  the  orifice  of  the  sense  organ  near  the  base  ; 
from  the  base  of  the  third  segment  also  arises  the  three-jointed  arista,  the  first  two 

segments  being,  however,  minute ;  the 
third  bears  a  series  of  from  seventeen 
to  twenty-one  fine  branched  hairs  on  one 
side.  The  male  genitalia  or  hypopygium 
is  more  or  less  oval  and  tumid,  its  long 
axis  lying  in  the  antero-posterior  direction, 
with  a  vulviform  median  groove  (the 
anus)  running  from  the  anterior  margin 
to  beyond  the  middle. 

[Newstead  has  shown  the  importance 
of  the  study  of  the  genitalia  in  separating 
species  {vide  Bull.  Ent.  Res.,  ii,  pp.  9-36 
and    107-110,  and   iii,  pp.   355-360;    and 
Anji.   Trop.  Med.   and  Par.,  vii,  No.    2, 
PP-  331-334)- 
[The  tsetse-flies  reproduce  differently  from  all  other  Miiscidce.     The  female  pro- 
duces at  each  birth  a  single  full-grown  larva,  which  is  retained  within  the  oviduct  and 
there  nourished  by  the  secretion  of  special  glands,  and  on  being  born  crawls  to  some 
hiding  place  and  at  once  becomes  a  puparium. 


Fig.  417. — Head   of    Glossina   longipalpis, 
Wied.     (After  Griinberg.) 


Fig.  418. — Antenna  of  Glossina pallidipes,  male.     (After  Austen.) 


[The  larva  is  a  yellowish  footless  maggot  nearly  as  large  as  the  mother's  body, 
the  skin  shagreened  and  the  anal  extremity  having  a  pair  of  large,  black,  granular 
prominences  separated  by  a  depression  containing  the  breathing  pores. 

[The  puparium  is  brown  of  various  shades,  the  tumid  lips  of  the  larva  being  con- 
spicuous,^the  size  and  shape  of  the  lips  enabling  the  puparia  to  be  identified. 


SPECIES   OF   GLOSSTNA 


605 


[These  puparia  are  often  found  in  masses  at  the  base  of  trees,  in  hollows  in  trees 
and  rocks  just  buried  under  vegetal  debris.  These  insects  are  generally  confined 
to  definite  tracts  known  as  "  fly-belts."  They  usually  occur  in  damp,  hot  places  on 
the  borders  of  rivers  and  lakes,  and  never  far  from  water  in  the  case  of  the  palpalis 
group,  although  others  of  the  morsitans  group  may  be  found  a  considerable  distance 
from  water.  They  are  usually  absent  on  grass  plains,  but  may  now  and  then  occur 
there  (Kinghorn,  vide  Hindles'  "  Flies  and  Disease,  Blood-sucking  Flies,"  1914, 
p.  274)  ;  cover  of  trees,  shrubs,  or  thick  reeds  is  essential  to  them. 

[Their  range  in  Africa  extends  roughly  from  18°  N.  to  31°  S. 

Sfilossina  palpalis  is  the  chief  carrier  of  the  more  prevalent  type  of  sleeping 
sickness.  Two  distinct  types  of  parasites  can  produce  this  disease,  viz.^  Trypano- 
soma gambiense^  which  produces  the  ordinary  sleeping  sickness,  transmitted  by 
G.  palpalis,  and  Trypanosoma  rhodesiettse  the  Rhodesian  or  Nyasaland  sleeping 
sickness,  transmitted  by  G.  7norsitans,  and  possibly  identical  with  T  brucei^  the 
parasite  of  N'agana.  Koch  has  also  shown  that  G.  pallidipes^  Austen,  and  G.fiisca^ 
Walker,  can  be  artificially  infected  with  the  human  trypanosome.  It  appears 
probable  that  Koch  used  G.  brevipaipis^  not  G.  fiisca^  in  his  transmission  experi- 
ments, as  at  that  \\m^  fusca  included  nearly  all  the  large  tsetses,  but  brevipalpis 
is  its  Eastern  representative. 

[A  Table  of  Species  (modified  after  Austen)  is  appended  here  :  — 


tachinoides,  West  wood. 


I. 

Glossifta  palpalis  GROUP. 

Dorsum  of  abdomen  ochraceous  buff  or  buff ;  third 

and  following  segments  exhibiting  sharply  defined, 

dark  brown  or  clove  brown,  interrupted  transverse 

bands    ... 
Dorsum  of  abdomen  not  so  marked  ...         ...         ...     2. 

Third  joint  of  antenntc  pale  (cream  buff  to  ochraceous 

buff),    clothed  with   long  and  fine  hair,  forming 

a  conspicuous  fringe  on  front  and  hind  margins        pallicera^  Bigot. 
Third  joint  of  antennas  entirely  dark  (mouse-grey) 

except  at  extreme  base  on  outer  side,  and  without 

a  conspicuous  fringe  of  long  and  fine  hair 
Dorsal    surface    of    abdomen    dark    sepia    brown  , 

median  paler  area  on  second  segment  broad,  and 

more    or  less  quadrate    or   irregular   in  outline 

hypopygium  of  $  buff  or  ochraceous  buff 
Dorsal  surface  of  abdomen  blackish-brown  ;  median 

paler   area  cuneate  {i.e.^   triangular   in  outline) 

hypopygium  of  $  grey        


3- 


caliginea^  Austen. 


palpalis^  Rob.  Desv. 


II. 

Glossi?ia  morsitans  Group. 

Hind  tarsi  entirely  dark  ;  small  slender  species  ; 
abdomen  bright  ochreous  or  reddish  ochreous 
with  dark  lateral  markings  

Hind  tarsi  not  entirely  dark  ;  abdomen  drab-grey, 
buff  or  ochreous  buff  with  conspicuous  dark 
interrupted  transverse  bands         


ausfenii,  Newstead. 


2. 


6o6 


THE   ANIMAL   PARASITES   OF   MAN 


Last  two  joints  of  front  and  middle  tarsi  with 
sharply  defined  clove  brown  or  black  tips 

Last  two  joints  of  front  and  middle  tarsi  without 
sharply  defined  clove  brown  or  black  tips  (front 
and  middle  tarsi  either  entirely  pale  or,  at  most, 
last  two  joints  of  front  tarsi  faintly  brownish  at 
the  tips),  and  last  joint  and  distal  half  of  penul- 
timate joint  of  middle  tarsi  light  brown,  never  so 
dark  as  to  form  a  sharp  contrast  with  the 
remaining"  joints 

Third  joint  of  antennas  with  a  distinct  fringe  of  fine 
hair  on  front  margin  ;  dark  brown  or  clove-brown 
bands  on  abdommal  segments  extendmg  close  to 
hind  margins  (z>.,  pale  ground  colour,  apart  from 
the  median  interspace,  confined  to  a  very  narrow 
hind  border) 

Third  joint  of  antennae  without  a  distinct  fringe  of 
fine  hair  on  front  margin  ;  dark  brown  or  clove- 
brown  bands  on  abdominal  segments  not  extend- 
ing close  to  hind  margins 


pallidipes,  Austen. 


lo?jgipalpis,  Wiedeman. 


morsitaiis^  Westwood 


in. 

Glossina  fusca  GROUP. 

Third  joint  of  antennae  fringed  with  fine  hair  on 
anterior  and  posterior  margins  ;  fringe  on  anterior 
margin  conspicuous  under  a  hand  lens  magnify- 
ing 1 5  diameters  (nominal)  when  head  is  viewed 
in  profile  ...  

Third  joint  of  antennae  with  fringe  of  fine  hair  on 
anterior  margin  so  short  as  to  be  scarcely  notice- 
able under  a  hand  lens  magnifying  15  diameters 
(nominal)  when  head  is  viewed  in  profile  (longest 
hairs  m  fringe  in  length  not  exceeding  one-sixth 
of  width  of  third  joint) ;  palpi  long  and  slender  ... 

Longest  hairs  in  fringe  on  front  margin  of  third 
joint  of  antennas,  in  length  equal  to  from  one- 
fourth  to  one-third  (not  exceeding  one-third)  of 
width  of  third  joint ;  palpi  of  moderate  length    ... 

Longest  hairs  infringe  on  front  margin  of  third  joint 
of  antennas  in  length  equal  to  from  one-half  to 
three-fourths  of  width  of  third  joint  ;  palpi 
noticeably  long  and  slender 

Pleurae  drab-grey  or  isabella-coloured,  hind  cox^ 
bufif  or  greyish-buff  .... 

Pleurae  dark  grey ;  hind  coxae  mouse-grey 


3- 


tabaniformis^  Westwood. 


nigrofusca^  Newstead. 

fusca.  Walker. 
fuscipleuris,  Austen. 


IV. 

Glossina  brevipalpis  GROUP. 

I.  Dorsum  of  thorax  with  four  sharply  defined  brown, 
more  or  less  oval  or  elongate  spots,  arranged  in 
a  parallelogram,  two  in  front  and  two  behind  the 


GLOSSINA   PALPALIS 


607 


transverse  suture  ;  proboscis  bulb  with  a  sharply 
defined  brown  or  dark  brown  tip   ... 

Dorsum  of  thorax  without  such  spots  ;  proboscis 
bulb  not  brown  or  dark  brown  at  tip 

Wings  with  upper  thickened  portion  of  anterior 
transverse  vein  much  darker  in  colour  than 
adjacent  veins  and  thus  standing  out  con- 
spicuously against  the  rest  of  the  wing     ... 

Wings  with  upper,  thickened  portion  of  anterior 
transverse  vein  not  much  darker  in  colour  than 
adjacent  veins,  and  thus  not  standing  out  con- 
spicuously against  the  rest  of  the  wings  (wings 
practically  unicolorous) 


longipennis,  Corti. 


brevipalpis^  Newstead. 


jnedicorum^  Austen. 


Glossina  palpalis,  Rob.  Desv. 

[This  is  the  chief  carrier  of  sleeping  sickness  in  Nature.  It  is  found  in  places 
over  the  whole  of  West  x^frica  from  the  mouth  of  the  Senegal  River  to  Angola, 
and  extends  eastwards  into  the  Bahr-el-Ghazal.  The  eastern  boundary  follows  the 
valley  of  the  Nile  and  includes  the  eastern 
shores  of  Lakes  Victoria  and  Tanganyika  ; 
from  the  southern  end  of  the  lake  the 
boundary  tends  south-west,  approximately 
following  the  frontier  between  North-eastern 
Rhodesia  and  the  Congo  Free  State,  and 
passing  through  the  Katanga  district  of 
the  latter  country  into  Angola  (Austen). 
It  may  occur  up  to  3,000  ft.;  but,  accord- 
ing to  Bagshawe,  it  has  not  been  recorded 
above  4,000  ft.  It  feeds  on  the  blood  of 
many  animals,  including  reptiles,  amphibia, 
birds,  and  even  amphibious  fishes,  as  well 
as  all  the  wild  mammals.  It  seems,  how- 
ever, to  possess  a  decided  predilection  for 
man,  and  undoubtedly  thrives  better  upon 
mammals  and  birds  than  upon  cold-blooded 
animals. 

[It  is  not  usually  found  far  from  water, 
requiring  a  humid  atmosphere  and  tem- 
perature of  about  85°  F.  (shade).  But  a 
marked  seasonal  distribution  is  shown,  the 
flies    considerably    extending    their    range 

during  the  rainy  season,  and  thus  visiting  districts  which  are  dry  for  the  greater 
part  of  the  year;  as  the  rains  diminish  the  fly  gradually  leaves  the  temporary 
haunts  and  returns  to  the  more  permanent  ones.  It  bites  only  by  day,  and  then 
only  in  sunny  weather,  and  usually  lives  in  shade. 

[Roubaud  has  shown  that  the  first  larva  produced  is  about  three  weeks  after 
copulation,  and  that  others  are  produced  at  an  interval  of  nine  or  ten  days.  The 
puparium    stage  is   rapidly   produced    after   the    expulsion    of    the   larva,  often    in 


Fig.  419. —  Glossina  palpalis  and   pupa 
rium.     (After  Brumpt.) 


'  Newstead  has  recently  described  another  species  as  G.  severini  {Ann.   Trap.  Med.  and 
Par.,  19 1 3,  vii,  No.  2,  pp.  331-334).     It  is  allied  to  G.  fuscipleuris,  Aust  n. 


6o8 


THE   ANIMAL   PARASITES   OF    MAN 


three-quarters  of  an  hour.  The  puparium  stage  lasts  from  thirty-two  to  thirty-five 
days.  The  puparia  occur  in  well-drained  humus  close  to  water,  sheltered  by  trees 
or  bushes,  in  crevices  in  rocks,  and  between  the  exposed  roots  of  trees,  sometimes 

in  sand. 

[Bruce  has  shown  that  only  a  very  small  percentage  of  flies  fed  experimentally 
on  infected  animals  ultimately  become  infective,  and  that  the  infectivity  of  this 
small  percentage  depends  upon  a  delayed  infection  of  the  salivary  glands. 

[A  variety,  wellmani  of  Austen,  is  found  in  Angola,  Gambia,  the  Katanga  district 
of  the  Congo  Free  State,  the  Matondwi  Islands  of  Tanganyika,  etc. 

Glossina    morsitans,  Westwood. 

[This  species  has  been  shown  by  Kinghorn  and  Yorke,  and  also  by  Bruce,  to  be 
responsible  for  the  transmission  of  Trypanosoma  rhodesiense,  the  micro-organism 
producing  sleeping  sickness  in  man  in  Rhodesia  and  Nyasaland  and  also  in  parts 
of  German  and   Portuguese    East   Africa.     Fisher  and   Taute   have  demonstrated 


Fig.  420. — The  tseise-fly  {Glossina  viorsilans,  Westwood). 


experimentally  that  Trypatiosoma  gainbiense — the  sleeping  sickness  parasite  of  other 
parts  of  Africa— may  also  be  transmitted  by  this  fly,  and  in  addition  it  is  known  to 
be  capable  of  disseminating  several  species  of  trypanosomes  pathogenic  to  animals. 
Of  these,  T.  brucei  {=  ?  T.  rhodesie7tse),  the  parasite  of  tsetse  disease,  first 
incriminated  by  Bruce,  is  perhaps  the  most  important. 

[It  is  the  most  widely  spread  of  all  tsetse-flies  ;  its  range  extends  from  Sene- 
gambia  in  the  north-west  to  Southern  Kordofan  and  Southern  Abyssinia  in  the 
north-east,  and  then  southwards  to  the  Bechuanaland  Protectorate,  North-eastern 
Transvaal  and  Zululand.  The  actual  localities  given  by  Austen  are  Gambia,  French 
Guinea,  Gold  Coast,  Togoland,  Dahomey,  Northern  Nigeria,  Congo  Free  State,  the 
Bahr-el-Ghazal,  the  Uganda  Protectorate,  German  East  Africa,  and  Portuguese  East 
Africa. 

[This  species  is  confined  to  "  belts,"  often  of  very  limited  extent,  and  appears 
to  prefer  regions  where  there  is  sufficient  vegetation  for  moderate  but  not  excessive 
cover  and  a   hot,   moderately  dry  climate.     It    is    not.  nearly   so   dependent    upon 


STOMOXYS  6C9 

water  as  is  G.'palpalis^  and  generally  is  most  active  in  a  dry  atmosphere  ;  some 
observers,  however,  state  that  in  certain  districts  it  is  more  common  along  the 
banks  and  edges  of  rivers.  This  tsetse-fly  has  been  taken  as  high  as  5,500  ft. 
altitude.  It  infests  native  villages  as  well  as  the  bush.  Like  other  tsetse-flies 
it  bites  not  only  during  the  hottest  part  of  the  day,  but  also  on  bright  warm  moon- 
light nights,  and  it  feeds  on  the  blood  of  all  mammals. 

[The  structure  of  the  male  genitalia  of  those  representatives  of  G.  inorsitans 
occurring  on  the  West  Coast  of  Africa  and  in  parts  of  the  Soudan  presents  certain 
constant  differences  from  that  of  the  typical  form  of  this  species  ;  this  form  is 
known  as  G,  morsitajis^  race  sub?norsitans,  Newst. 

Genus.     Stomoxys,  Geoffroy. 

[The  members  of  this  genus  which  occur  in  temperate  and  tropical  countries 
are  provided  with  a  hard,  slender,  shiny  black  proboscis  which  projects  horizontally 
from  beneath  the  head ;  by  means  of  this  structure  they  can  bite  severely. 
In    general    appearance    they    resemble    house   flies,    but    the    proboscis   at   once 


Fig.  421. — The  stinging  fly  {Stomoxys  calcitrans,  Linn.). 

distinguishes  them.  In  many  parts  of  Britain  they  are  known  as  storm  flies 
on  account  of  their  frequent  appearance  indoors  previous  to  a  storm  of  rain  or 
wind,  which  I  have  invariably  found  to  be  correct ;  they  are  also  called  stinging 
flies.  In  colour  they  are  greyish,  dusky  or  brownish-grey  or  black,  varying  from 
5  to  7  mm.  in  length;  the  thorax  has  dark  longitudinal  stripes  and  the  abdomen 
dark  spots  or  bands.  In  the  male  the  eyes  are  closer  together  than  in  the  female. 
These  flies  usually  occur  in  stables  and  farmyards,  along  woods  and  in  lanes,  and 
mainly  attack  mammals. 

[One  s^G:c\Qs{Slo?noxys  calcitrans,  Linnaeus)  occurs  practically  all  over  the  world. 
The  female  lays  her  eggs  in  moist,  warm,  decaying  vegetation  ;  as  many  as  eighty 
may  be  laid  by  a  single  female.  The  ova  are  white,  banana-shaped,  with  a  broad 
groove  on  the  shorter  curvature  ;  they  may  hatch  in  two  or  three  days.  The 
creamy-white  larva  tapers  to  a  point  at  the  head  end,  and  is  truncated  at  the  tail 
end.  Two  black  mouth  hooks  are  plainly  visible  at  the  cephalic  extremity.  There 
are  two  plates  on  the  posterior  surface  of  the  last  segment  which  bear  the  respiratory 
pores,  nearly  circular  in  outline.  It  reaches  maturity  in  fourteen  to  twenty-one 
days;  when  mature  it  is  11  mm.  long.     The  pupal  stage  is  passed  in  the  old  larva 


6io  THE   ANIMAL    PARASITES   OF   MAN 

skin  and  lasts  from  nine  to  thirteen  days  ;  it  is  barrel-shaped,  5  to  8  mm.  long,  and 
of  a  bright  reddish-brown  to  dark  chestnut-brown  colour. 

[This  insect  may  act  as  a  carrier  of  anthrax,  and  has  been  proved  to  be  the  agent 
of  an  extensive  epidemic  of  malignant  pustule  in  the  Isle  of  Pines,  New  Caledonia.^ 

[Noe's-  experiments  tend  to  show  that  it  is  an  intermediate  host  and  transmitter 
of  Filaria  labiato-papillosa  of  the  ox. 

[Surra  is  generally  stated  to  be  transmitted  by  Stomoxys  as  well  as  Tabanus, 
and  yet  Nitzman  in  the  Philippines  obtained  uniformly  negative  results  in  exhaustive 
experiments.  Others  have  also  been  unsuccessful.  Certainly  Stomoxys  can  transmit 
the  disease  in  French  West  Africa  (Bonet  and  Roubaud),  and  mechanically  has 
been  proved  to  be  capable  of  disseminating  other  trypanosomes  (experimentally)  : 
sleeping  sickness  (J",  gcunbiense) ;  nagana  (7".  brucei) ;  sou  ma  (7".  cazalboui)  ;  and 
el  debat  (Z'.  soudane7ise). 

[5.  calcitrajis  may  also  be  a  carrier  of  poliomyelitis  (Rosenau  and  Brues, 
Harvard  Alumni  Bulletin,  191 2,  xv,  No.  9,  pp.  140-142).  Several  species  are 
ow  known  (5.  brunnipes^  Griinb. ;  S.  inornata,  Griinb. ;  S.  fiigra^  Macq.  ; 
S.  omega^   Newst.  ;   S.  ochrosoma^   Speiser,  etc.). 

Genus.     Lyperosia,  Rondani. 

[A  genus  of  small  flies  which  bite  man  and  animals,  but  are  not  so  far  connected 
with  the  transmission  of  any  disease  in  man,  but  in  Java  it  appears  to  carry  surra 
(P.  Schat,  Meeledeel  Praefstatioti  Oost-Java,  1903,  3e  ser..  No.  44),  the  species  being 
Lyperosia  exigua,  Meijere.  These  flies  can  be  told  from  Stomoxys  by  the  palpi 
being  broader,  flattened  laterally,  and  as  long,  or  nearly  so,  as  the  proboscis. 
When  not  feeding  the  palpi  enclose  the  proboscis,  as  in  Glossina.  They  are  usually 
about  half  the  size  of  Stomoxys,  and  are  the  smallest  blood-sucking  Muscidcc. 
They  frequently  swarm  around  and  upon  domesticated  animals. 

[The  life-history  of  the  horn  fly  in  America  (Z.  irritans,  Linn.)  is  well  known. 
It  lays  its  ova  singly  in  freshly  dropped  cow-dung,  and  there  the  maggots  feed, 
pupating  in  the  soil  beneath. 

[Patton  and  Cragg  also  give  some  details  as  to  the  life-history  of  Liperosia 
exigua  ("Medical  Entomology,"  p.  375)  as  follows:  "  Z.  exigiia,  whose  habits 
have  been  observed  in  Madras,  usually  lays  twelve  eggs  at  a  time.  The  flies 
immediately  return  to  the  cow  and  the  process  is  repeated  when  the  dung  is  again 
dropped.  The  larvae  migrate  from  the  dung  when  about  to  pupate,  and  the  puparia 
are  always  found  in  the  earth  at  some  distance  away  or  under  the  sides  of  the  patch 
of  dung.  The  fly  usually  hatches  out  in  five  days,  though  sometimes  as  late  as  the 
eighth.  Weiss  has  studied  the  life-history  of  irrila?ts  var.  weisii  from  Algeria  ; 
its  larval  stage  lasts  five  days,  and  the  flies  hatch  out  of  the  puparia  in  another 
five  days." 

[The  other  biting  genera  oi  Muscidce,  Hsematobia,  Hasmatobosca,  Bdellolarynx, 
Stygeromyia,  and  Philaematomyia,  although  sometimes  annoying  to  man,  have 
not  in  any  way  been  connected  with  any  disease. 

[The  horse  fly  {Hcnnatobia  irrita?is,  L."^)  attacks  cattle  chiefly,  but  now  and  then 
man  is  bitten.  The  different  species  can  be  told  from  Stomoxys  by  the  palpi  being 
nearly  as  long  as  the  proboscis. 


'  Bull,  des  Seances  de  la  Soc.  ent.  de  France,  1878,  pp.  cxliv,  cxlv. 

==  Atti  della  Reale  Accad.  del  Lincei,  Anno  CCC.   Se  Quinta,   1903,  xii,   2  sem.   fasc, 
PP-  387-393- 

2  This  is  apparently  the  stimulans  of  Meigen. 


PUPIPARA  6ll 

[The  genus  Philaematomyla,  Austen,  is  intermediate  between  Stomoxys  and 
Musca  in  structure,  and  between  the  non-blood-sucking  Musca,  as  M.  domestica^ 
and  the  blood-sucking  Musca  pattoni^  Austen,  which  feeds  on  the  blood  exuding 
from  the  bites  of  true  blood-suckers.  They  occur  in  Central  Africa  and  India, 
Ceylon  and  Cyprus  {vide  "  The  Life-history  of  PhilcEmatomyia  insignis^  Austen," 
Ann.  Trap.  Med.  and  Par. ^  191 2,  v,  p.  515). 

[Two  flies  belonging  to  the  family  Anthomyidae  also  attack  man,  namely  : — 

\HydrotcEa  jneteorica,  L.  (the  meteoric  fly).  This  fly  attacks  man  as  well  as 
animals.  They  especially  bite  around  the  eyes  and  nostrils  of  animals,  but  are  not 
so  particular  with  man  ;  the  head,  however,  is  usually  chosen.  Linnaeus  called  it 
the  meteoric  fly  because  it  often  forms  clouds  around  horses'  heads  at  the  approach 
of  rain.  The  Hydrotasas  are  usually  black  or  blue-black  in  colour  with  bare  eyes 
and  simple  abdomen,  the  front  femora  peculiarly  constructed.  H.  meteorica,  L., 
occurs  in  Britain. 

[The  members  of  the  genus  Hydrophoria,  Desvoidy,  also  bite  man. 

Pupipara  or  Eproboscidae. 

[The  Pupipara  are  all  blood-suckers,  the  majority  occurring  as  parasites  on 
mammals  and  birds,  where  they  are  more  or  less  permanent  parasites.  Occasionally 
some  may  attack  man.  They  all  produce  their  young  fully  formed,  and  they  assume 
the  pupal  stage  immediately  after  extrusion.  The  puparia  are  large.  They  are 
mostly  flat,  louse-like  flies  which  may  or  may  not  be  winged.  In  the  case  of  Melo- 
phagus  I  have  found  the  puparia  are  often  passed  by  the  female.  The  winged  forms 
have  a  short  quick  flight,  and  when  disturbed  will  seek  shelter  in  man's  hair  or 
beard.  Two  main  families  occur  :  (i)  the  HippoboscidcF,  and  (2)  the  NycteribiidcE. 
The  former  occur  on  animals  and  birds,  the  latter  on  bats  only,  but  may  invade 
man.  Two  other  families  are  known — the  Braulidce  (bee  parasites)  and  the  Streblidce 
(bat  parasites). 

[The  mouth  of  the  Hippoboscidce  is  long  and  sharp,  forming  a  proboscis.  The 
thorax  and  abdomen  are  flat  and  leathery.  The  legs  are  stout  and  strong,  and 
terminate  in  large  dentate  claws  and  other  structures  of  use  in  holding  on  to  the 
hair  or  feathers  of  their  host  when  blood-sucking. 

[Austen  says  it  is  probable  that  the  Hippoboscidce  are  descended  from  ancestors 
belonging  to  the  MuscidcB.,  which  underwent  modification  in  bodily  structure  as  the 
consequence  of  the  adoption  of  a  parasitic  mode  of  life. 

[Two  wings  are  present  in  the  true  Hippoboscse,  Hippoboscaequitia  (of  the  horse), 
H.  camelina  (of  the  camel),  H.  maculata  (of  oxen),  and  H.  capejtsis  (of  dogs),  but 
are  absent  in  Melophagus,  the  sheep  tick  or  ked  fly  {M.  oviniis). 

[In  two  genera,  Lipoptena  and  Echestypus,  wings  are  at  first  present,  but  are 
lost  as  soon  as  the  fly  finds  its  permanent  host. 

[With  regard  to  their  biting  man,  such  is  only  occasional.  I  have  known  sheep 
shearers  to  be  badly  bitten  by  Melophagus  ovinus,  and  have  more  than  once  been 
attacked  myself  when  standing  where  shearing  is  taking  place.  Sharp  records  the 
grouse  parasite,  O^^nithomyia  lagopodis^  as  once  biting  severely  a  gamekeeper  in 
Scotland.     There  are  also  records  of  H.  inaculata  biting  man  in  Africa  and  India. 

[Although  so  far  not  connected  with  any  human  disease,  it  is  interesting  to  note 
Theiler  has  shown  that  Hippobosca  rufipes^  v.  Olfers,  and  H.  maculata^  Leach,  are 
capable  of  transmitting  Trypanosoma  theileri^  Laveran,  the  cause  of  gall  sickness 
amongst  cattle  in  the  Transvaal.  It  is  now  considered,  however,  that  Trypano- 
soma theileri  is  non-pathogenic,  and  that  the  cause  of  gall  sickness  is  a  piro- 
plasma-like    organism    known    as   A7iaplasma    margi7iale.      Theiler,    Laveran    and 


6l2  THE   ANIMAL   PARASITES   OF   MAN 

Mesnil  all  hold  this  view  {vide  Laveran  and  Mesnil,  "  Trypanosomes  and  Trypano- 
somiases," second  edition,  191 2,  p.  330). 

[Zjw^-^m.— Three  members  of  this  genus  have  been  shown  to  transmit  the 
non-pathogenic  (?)  organism,  HceDioproteus  columbcE  amongst  pigeons  in  Algeria 
and  S.  America. 

Insects  and  Epidemic  Poliomyelitis. 

[In  a  recent  number  of  the  Joicrftal  of  Economic  Entomology}  Brues  and 
Sheppard  point  out  the  possibility  of  acute  epidemic  poliomyelitis  (infantile  paralysis) 
being  an  insect-borne  disease.     They  summarize  as  follows  : — 

[Many  facts  connected  with  the  distribution  of  cases  and  the  spread  of  epidemics 
of  this  disease  with  histories  of  insects  bites,  suggest  at  least  that  the  disease  may 
be  insect-borne.  Field  work  during  the  past  summer,  together  with  a  consideration 
of  the  epidemiology  of  the  disease  so  far  known,  points  strongly  towards  biting 
flies  as  possible  carriers  of  the  virus.  It  seems  probable  that  the  common  stable 
fly  {Stomoxys  calcitrmts^  L.)  may  be  responsible  to  a  certain  extent  for  the  spread 
of  acute  epidemic  poliomyelitis,  possibly  aided  by  other  biting  flies  such  as 
Tabanus  lineola.  No  facts  which  disprove  such  a  hypothesis  have  as  yet  been 
adduced,  and  experiments  based  upon  it  are  now  in  progress. 

[If  the  disease  should  prove  to  be  common  to  any  species  of  domestic  animals, 
as  is  now  strongly  suspected,  a  secondary  connection  of  ticks  in  spreading  the 
disease  among  such  animals  seems  probable,  as  has  been  mentioned. 

[The  following  is  some  of  the  more  important  literature  on  Diptera  in  general :  Meigen, 
J.  W.,  "  Syst.  Besch.  d.  bek.  europ.  zweifliigligen  insecten,"  1818-1838,  7  vols.  ;  Brauer,  F.» 
"Monographic  der  Oestriden,"  Wien,  1863;  Idem,  "Nachtr.  hiersu,  "  Wien.  ent.  Zeit.i 
1887,  vi,  pp.  4,  71  ;  Schiner,  J.  R.,  "  Fauna  austriaca  :  die  Fliegen,"  Wien,  1860-64;  Low,  Fr., 
"  Ueber  Myiasis  und  ihre  Erzeuger,"  Wien.  vied.  Wochenschr.,  1882,  xxii,  p.  247;  1883, 
xxxiii,  p.  972;  Joseph,  G.,  "  Ueb.  Fliegen  als  Schadlinge  und  Parasiten  des  Menschen," 
Deiitsch.  vied.  Zeit.,  1885,  i,  p.  37  ;  1887,  iii,  pp.  713  and  725  ;  Peiper,  E.,  "  Fliegenlarven  als 
gelegentl.  Paras,  d.  Mensch.,"  Berlin,  1900;  Theobald,  F.  V.,  "Monograph  of  the  Culicidai 
of  the  World,"  1901-1911,  5  vols,  and  i  atlas,  plates  ;  Austen,  E.,  "A  Monograph  of  Glossina 
Tsetse-flies,"  1903,  i  vol. ;  Van  der  Wulp,  "  Diptera  neerlandica,"  1877  ;  Walker,  "  Insecta 
Britannica  :  Diptera,"  1851-53  and  1856;  Lundbeck,  "Diptera  danica,"  1907-12;  Zetterstedt, 
"Diptera  scandinavire,"  1850  ;  Theobald,  "British  Flies,"  1892;  Aldrich,  "  N.  American 
Diptera,"  1905  ;  Loew  and  Osten  Sacken,  "Monographs  of  the  N.  American  Diptera,'' 
1862-63  and  1869;  Macquart,  "Diptera  exotique,"  1830-47;  Rondani,  "Diptera  exotica  et 
Italica,"  1863-68  ;  Williston,  "  Manual  of  Famiiies  and  Genera  of  N.  American  Diptera," 
second  edition  ;  Verrall,  "  British  Flies."  A  fuller  literature  will  be  found  in  Peiper,  as  well  as 
in  Ruber's  "  Bibliographie  d.  klin.  Ent.,"  1899,  iii,  Jena,  in  the  Bibliography  at  the  end  of 
this  work  and  in  the  Rev.  of  App.  Ent.  (Dulau  and  Co.,  London),  where  all  references  to 
modern  research  can  be  found. — F.  V.  T.] 


'   Charles  T.  Brues  and  Philip  A.  E.  Sheppard,  "The  Possible  Etiological  Relation  of 
certain  Biting  Insects  to  the  Spread  of  Infantile  Paralysis," /^«r;/.   Econ.  E71L,  1912,  cciv, 


pp.  305-324- 


ADDENDA  613 


ADDENDA. 

Akamushi  or  Kedani  Sickness  (vide  also  p.  487). — Schutfner  (Far 
East.  Assoc.  Trop.  Med.,  Conipi.  rend.  Trois.  Cong.  Biennial^  19^3, 
Saigon,  1914,  pp.  309-315)  states  he  observed  a  peculiar  fever  in  Deli, 
Sumatra,  somewhat  resembling  typhoid.  This  he  traced  either  to  a 
mite  or  tick.  He  figures  the  possible  carriers,  namely,  a  Trombidium 
and  Cheyletida'.  He  calls  this  disease  pseudo-typhus — a  variant  of 
Japanese  kedani  sickness,  which,  he  says,  also  occurs  in  the  Philippines. 

Ticks. — African  Tick  Fever  :  Marzinovsky  (Proc.  of  Conference  of 
Bacteriologisfs  and  Representatives  of  Medical  Sanitary  Authorities  on 
the  Campaign  against  Infections  Diseases  in  connection  with  the  War, 
Soc.  Rnss.  Physicians  in  mem.  Pirosov,  Moscow,  191 5,  pp.  56-68),  states 
that  African  tick  fever  has  been  imported  into  Persia,  and  that  it  is 
there  carried  by  Ornithodorus  tholosani. 

Tick  Pakalysis  :  Todd  ("  Paralysis  and  Tick-bite,"  Can.  Med. 
Assoc.  Jonrn.,  1914,  iv.  No.  9,  pp.  825-826)  refers  to  paralysis  ascribed 
to  the  bites  of  ticks  in  children,  and  possibly  adults,  in  America, 
British  Columbia  and  Australia.  He  states  that  a  young  child, 
perfectly  well  one  day,  has  more  or  less  complete  paresis  or  paralysis 
on  the  next,  fever,  a  rapid  pulse,  and  other  constitutional  symptoms. 
The  child  may  be  dull  and  stupid,  and  may  have  convulsions.  If 
the  tick  is  not  found  and  removed  the  child  may  die,  but  if  it  is 
removed,  the  symptoms  disappear  and  recovery  is  complete  in  a 
few  hours.      The  tick  must  be  entirely  removed. 

Diptera. — Psychodid^  :  Bolt  {China  Med.  Jonrn.,  Shanghai, 
xxix.  No.  2,  pp.  78-86)  states  that  sand-flies  (Plebotomus)  and  the 
fever  due  to  them  are  common  m  North  China,  May  and  June  being 
the  worst  months.  The  natives  of  the  region  appear  to  be  immune, 
but  all  others  suffer.  Old  ruined  buildings  are  the  favourite  haunts 
of  the  Phlebotomus.  The  species  of  Phlebotomus  has  not  been 
determined. 

Pulicidae. — Dermatophilus  (Sarcopsylla)  penetrans,  or  the 
*' Jigger." — This  flea  (r/t/d  p.  544)  is  believed  by  Lama  {Giorn.  Ital. 
Mai.  Yen.,  Milan,  1914,  xlix,  pp.  465-472)  frequently  to  carry  leprosy 
and  he  points  out  that  the  early  lesions  of  leprosy  usually  appear 
on  the  uncovered  parts  of  the  body.     This  flea  also  attacks  rats. 

Brachycera.  — Leptid^  {vide  p.  603) :  White,  A.  {''  The  Diptera- 


6l4  THE   ANIMAL   PARASITES   OF   MAN 

Brachycera  of  Tasmania,"  part  I,  Papers  and  Proc.  Roy.  Soc.  of 
Tasuiaiiia  for  1914,  191 5,  pp.  35-74),  describes  a  new  blood-sucking 
Leptid;  Spaniopsis  tahaniformis,  which  resembles  a  small  gad  fly 
(Tabanus)  in  appearance. 

Pycnosoma  putorium :  This  is  believed  by  Roubaud  ("  Les  Pro- 
ducteiirs  de  Myiases  et  Agents  similaires  chez  I'homme  et  les  ani- 
maux,"  Paris,  191^,  part  I)  to  be  largely  concerned  in  the  spread  of 
amoebic  dysentery  in  French  West  Africa. 

LtLciUa  argyrocephala,  Macquart :  This  green-bottle  fly  is  described 
by  Roubaud  as  producing  myiasis  in  Africa  ("  Les  Producteurs  de 
Myiases  et  Agents  similaires  chez  I'homme  et  les  animaux,"  19 14, 
Paris,  part  I).     It  attacks  ulcers  and  sores  in  man  and  animals. 

Aiichmeromyia  liiteola,  Fabr.  :  Schwetz  (Ann.  Trop.  Med.  and  Par., 
19 14,  viii.  No.  3,  pp.  497-507),  collected  a  large  quantity  of  this 
insect  at  Kabinda.  He  placed  them  in  flasks  with  sand  and  a  few 
days  later  they  pupated,  and  in  fifteen  days  several  flies  hatched 
out.  The  larval  period  varies  from  an  unknown  minimum  up  to 
several  months.  The  larva  may  live  for  at  least  two  months  without 
food.  A  female  oviposited  on  the  17th,  and  on  the  i8th  one  larva 
hatched.  The  pupal  stagei  seems  to  last  eight  to  fifteen  days.  The 
larvae  appear  to  bite  by  day  as  well  as  night  according  to  native 
m  format  ion. 

Cordylobia  anthropophaga,  Griinb.  :  Roubaud  ("  Etudes  sur  la 
Faune  parasitaire  de  I'Afrique  occidentale  frangaise,"  part  I,  ''Les 
Producteurs  des  Myiases  et  Agents  similaires  chez  I'homme  et  les 
animaux,"  Paris,  1914)  gives  the  life-history  of  this  species.  One 
fly  laid  150  ova  in  a  glass  vessel,  on  the  sides,  and  on  some  rotten 
fruit,  and  died  the  following  day.  He  found  that  fifteen  larva?  just 
hatched  placed  on  sand  in  a  glass  vessel  with  a  guinea-pig  gave  rise 
to  characteristic  tumours  on  the  ventral  surface  of  the  body  and 
the  anus.  Other  experiments  failed.  It  thus  seems  that  infection 
takes  place  from  larva?  which  have  hatched  apart  from  the  host. 
Infection  of  man  is  regarded  as  accidental;  no  positive  infection  of 
horses,  oxen,  sheep  or  pigs  is  known— it  is  rare  in  goats,  and  poultry 
never  seem  to  be  attacked.  The  result  of  experiments  tends  to  show 
that  the  apparent  choice  of  a  host  is  mainly  a  question  of  body 
temperature.  The  larva,  whether  freshly  emerged  or  eight  to  ten  days 
old,  penetrates  the  skin  immediately,  boring  obliquely  between  the 
epidermis  and  dermis.  Once  removed  from  the  tumour  the  maggot 
cannot  bore  again.  The  first  moult  takes  place  about  three  days 
after  penetration,  and  the  total  period  of  residence  in  the  host  is 
seven  to  eight  days.  Upon  emerging  the  larva  falls  to  the  ground 
and  buries  itself.  In  two  or  three  days  it  pupates  and  this  stage  lasts 
no  longer  than  twenty  days.  High  temperatures,  such  as  95°  F., 
appear  to  be  fatal. 


ADDENDA  615 

Myiasis.— Coates,  G.  M.,  "  A  Case  of  Myiasis  Aurium  accom- 
panying the  Radical  Mastoid  Operation,"  Jotirn.  Amer.  Med.  Assoc, 
Chicago,  111.,  19 14,  Ixiii,  pp.  479-480  :  Apparently  C.  macellaria,  forty 
to  fifty  coming  away  with  the  gauze  after  the  operation. 

Huber,  G.  U.,  and  Flack,  F.  L.,  "An  Unusual  Case  of  Screw- 
worms  in  the  Nose  and  Nasal  Accessory  Sinuses/'  Journ.  Amer.  Med, 
Assoc,  Chicago,  1914,  Ixiii,  pp.  2288-2289. 

Auricular  Myiasis. — Francaviglia,  M.  C,  "An  cora  sulla  myiasi 
auricolare,"  Boll.  Sedute  Accad.  Gioenia,  Catania,  1914,  No.  31,  pp. 
15-23.  This  writer  mentions  the  following  parasites  in  the  human  ear  : 
Sarcophaga  carnaria,  L. ;  Wohlfartia  uiagiiifica,  Schiner ;  Chrysomyia 
macellaria,  F. ;  CalUphora  vomitoria,  L. ;  and  Anthomyia  pliivialis,  L. 
He  refers  to  a  severe  myiasis  in  Russia,  due  to  a  fly  variously 
recorded  as  Sarcophaga  zvohlfarti,  Rond. ;  S.  ruralis,  Meig. ;  or  Sarco- 
pliila  meigeni,  Portsch.  These  are  all  probably  synonyms  of 
W.  magnifica.  Chrysomyia  macellaria,  in  Central  America  and 
South  America,  is  quite  as  harmful  as  S.  carnaria,  causing  perfora- 
tion of  the  tympanum  and  meningitis.  Lucilia  nobilis  and  L.  ccesar 
have  also  been  incriminated.  Of  the  sub-family  Anthomyince,  the 
larvae  of  Fannia  scalaris,  Meig.,  F.  canicidaris,  Meig.,  F.  incisiirata, 
Zett,  and  Hydrotcea  meteorica,  L.,  are  chiefly  associated  with 
myiasis.  He  recommends,  if  the  larvae  are  outside  the  tympanum, 
an  injection  of  chloroform  vapour  by  a  few  drops  of  water  saturated 
with  chloroform,  by  an  emulsion  of  5  per  cent,  carbon  bisulphide 
or  with  benzine.  When  detached  they  may  be  removed  with  forceps 
or  a  solution  of  boric  acid.  If  the  tympanum  has  been  perforated, 
the  larvae  must  be  removed  at  once. 

Francaviglia  also  records  the  larva  of  Oestrus  ovis  in  the  human 
ear  {Boll.  Sedute  Accad.  Gioenia,  Catania,  1914,  No.  31,  pp.  23-27). 

Body,  Head,  and  Clothes  Lice.  —  Lobaczewski  (IVien.  klin. 
Wochenschr.,  Vienna,  191 5,  xxviii,  pp.  373-374)  recommends  the 
impregnation  of  body  linen  with  a  30  per  cent,  solution  of  oleum  betas 
in  96  per  cent,  alcohol  as  an  efficient  method  of  keeping  the  body 
free  of  lice.  But  the  process  must  be  renewed  each  time  the  linen  is 
washed  and  it  takes  fifteen  minutes  to  carry  out.  On  adding  the 
oil  to  the  alcohol,  a  portion  of  the  former  is  precipitated,  the  super- 
natant fluid  is  decanted  and  poured  over  the  linen,  which  is  wrung 
out  in  it  and  dried.  The  garments  retain  their  lice-proof  properties 
until  washed.  Three  days  after  wearing  the  clothes  thus  treated  no 
lice  remain  on  the  body. 

Portnikov,  Proc.  of  Conference  of  Bacteriologists  and  Representatives 
of  Medical  Sanitary  Author ities  on  the  Campaign  against  Infectious 
Diseases  in  connection  with  the  War,  Soc  Russ.  Physicians  in  mem. 
Pirosov,  Moscow,  191 5,  p.  131). 


6  [6  THE    ANIMAL    PARASITES   OF   MAN 

Pediculus  capitis  and  Plithinis  pubis  are  shown  to  be  successfully 
controlled  by  applying  spirit  extract  of  sabadilla  and  both  white  and 
grey  mercury  ointment,  solution  of  corrosive  sublimate  of  a  strength 
of  I  in  250  to  I  in  100,  amyl  and  ethyl  alcohol,  benzine,  chloroform, 
carbon  tetrachloride,  methane,  birch  tar,  liquid  of  malinin,  etc.  The 
control  of  Pediculus  vestiuieuti  by  the  mixture  of  tartaric  acid  and 
sodium  sulphite  slightly  moistened  with  water  is  advised.  It  is  placed 
HI  small  linen  bags  underneath  the  shirt ;  the  heat  of  the  body  produces 
a  reaction  which  continues  for  two  days,  giving  off  a  large  amount  of 
SO2,  which  spreads  beneath  the  shirt  and  kills  all  the  parasites  but  does 
not  affect  the  skin.  Marzinovsky,  in  the  same  Pioceedings  (pp.  56-68), 
gives  a  number  of  remedies  for  Pediculus  i^estiuienti  (called  liuuiauus), 
and  mentions  quinine  or  mercury,  which  latter  the  natives  in  Tur- 
kestan carry  on  their  hands  and  legs  in  bracelets  soaked  in  mercury 
compounds.  He  also  mentions  ethereal  oils,  the  most  effective  being 
clove  oil,  eucalyptus,  oil  of  anise  and  camphor.  He  recommends  for 
disinfecting  clothing  for  army  purposes  the  chamber  used  by  the 
Japanese  on  a  large  scale.  Kummerfelds'  wash  is  advised,  and  is  pre- 
pared as  follows :  20  parts  of  precipitated  sulphur  are  incorporated  in 
a  mortar  with  50  parts  of  glycerine  ;  2  parts  of  camphor  are  separately 
ground  with  50  of  eau-de-Cologne  and  20  of  borax,  and  870  parts  of 
distilled  water  are  added ;  the  whole  is  mixed  together  and  3  drops  of 
an  extract  of  musk  are  added  ;  shake  in  order  to  prevent  the  sulphur 
settling  down;  50  parts  of  ether  are  added  to  the  mixture.  This 
sounds  an  expensive  and  troublesome  preparation  to  make. 

Shipley  A.  E.,  '*  F'lowers  of  Sulphur  and  Lice,"  Brit.  Med.  fount., 
1915,  p.  295.  It  is  here  stated  by  Dr.  Lounsbury  that  the^  South 
African  troops  were  supplied  by  the  Government  with  bags  of  flowers 
of  sulphur  sewn  in  small  calico  bags  and  secured  to  the  underclothing 
next  the  skin  as  a  preventive  of  lice.  The  bags  were  2  in.  square, 
one  on  the  trunk  and  one  against  each  leg.  This  is  a  generally 
accepted  preventive,  but  is  best  mixed  with  equal  parts  of  creosote 
and  naphthalene. 

Shipley,  A.  E.,  ''  Insects  and  War,"  Brit.  Med.  Jouru.,  September 
19  to  November  14,  1914.     General  advice  given  re  lice. 


SUPPLEMENT  617. 


SUPPLEMENT: 

CLINICAL   AND    THERAPEUTICAL    NOTES. 

PROTOZOA. 

INTRODUCTION. 

The  aim  of  the  present  volume  is  to  give  an  account  of  the  animal 
parasites  of  man,  the  number  of  which  is  very  large.  The  Protozoa 
that  infest  man  are  very  important,  and  the  literature  relating  to  them 
and  to  the  treatment  of  the  diseases  that  they  produce  is  very  exten- 
sive. All  that  can  be  done  in  this  Appendix  is  to  give  a  very  brief 
outline  of  some  of  the  more  recent  and  approved  methods  of  treat- 
ment, for  further  details  of  which  the  reader  should  refer  to  standard 
medical  works,  among  which  the  following  are  noteworthy  : — 

AUbutt  and  Rolleston  (1907):  "System  of  Medicine,"  vol.  ii^ 
part  2,  "Tropical  Diseases  and  Animal  Parasites,"  London. 

Castellani  and  Chalmers  (191  3):  "Manual  of  Tropical  Medicine" 
(second  edition),  London. 

Laveran  and  Mesnil  (1912)  :  "  Trypanosomes  et  Trypanosomiases '" 
(second  edition),  Paris. 

Manson  (1914)  :  "Tropical  Diseases"  (fifth  edition),  London. 

Mense  (1905)  :    "  Handbuch  der  Tropenkrankheiten,"  Leipzig. 

Ross  (191 1 )  :    "  The  Prevention  of  Malaria,"  London. 

Scheube  (19 10)  :    "  Die  Krankheiten   der  Warmen  Lander,"  Jena. 

References  to  the  treatments  tried  in  many  parasitic  diseases  can 
be  found  in  the  Sleeping  Sickness  Bulletin  and  Kala-azar  Bulletin,  both 
now  superseded  and  greatly  extended  in  scope  in  the  Tropical  Diseases 
Bw/Zd/Z/f,  published  by  the  Tropical  Diseases  Bureau,  Imperial  Institute,. 
London,  S.W. 

The  following  diseases,  due  to  protozoa  and  allied  forms,  are. 
discussed  : — 

I.     Amoebic  Dysentery. 

II.    'Trypanosomiases. 

III.  Flagellate  Diarrhoea  and  Dysentery. 

IV.  Leishmaniases — Kala-azar  and  Oriental  Sore. 

V.  Spirochaetoses  —  Relapsing     Fevers,     Yaws,      Syphilis     and 

Bronchial. 

VI.  Malaria. 

VII.     Balantidian  or  Ciliate  Dysentery. 


6l8  THE   ANIMAL   PARASITES   OF   MAN 

I— AMCEBIC  DYSENTERY. 

Amoebic  dysentery,  due  to  Eniamceba  hisiolytica  (see  pp.  34-41), 
is  present  throughout  the  tropical  world  and  also  occurs  in  temperate 
zones. 

Walker  and  Sellards^  (1913)  conducted  important  experiments 
with  amoebae  on  prisoners  in  the  Philippine  Islands.  They  showed 
experimentally  that  cultural  amoebae  are  non-pathogenic.  As  regards 
experiments  w^ith  Entamoeba  coli,  after  feeding  to  twenty  individuals 
they  concluded  that  E.  coli  is  a  parasite  of  the  human  intestine  but 
non-pathogenic  and  non-culturable.  In  a  third  series  of  experiments, 
after  feeding  with  motile  Entamoeba  histolytica,  tetragena  cysts  were 
found  in  the  stools  later;  when  tetragena  cysts  were  administered, 
motile  E.  histolytica  were  present  in  the  subsequent  stools.  Some  of 
the  histolytica  cases  developed  dysentery  after  a  time.  They  lay  stress 
on  the  necessity  for  the  frequent  examination  of  stools  in  order  to 
detect  carriers.  The  incubation  period  of  entamcebic  dysentery  is 
usually  long. 

With  regard  to  the  symptomatology  of  amoebic  dysentery,  Castellan i 
and  Chalmers  distinguish  four  types — the  acute,  chronic,  latent,  and 
mixed  types. 

The  acute  type  has  an  abrupt  onset  ;  pain  is  felt  in  the  lower  part 
of  the  abdomen,  and  the  motions,  rarely  exceeding  thirty  daily,  are 
accompanied  by  much  griping  and  straining.  Blood  and  mucus  are 
present  in  the  motions,  and  occasionally  greyish  material,  consisting 
of  leucocytes,  mucus,  Charcot-Leyden  crystals,  amoebae,  and  bacteria, 
sometimes  with  particles  of  tissue.  Nausea  and  vomiting  may  occur, 
Digestion  is  usually  deranged.  The  abdomen  is  sunken,  the  liver  and 
spleen  are  normal,  but  tenderness  is  felt  along  the  course  of  the  large 
intestine.     The  urine  may  be  diminished  in  quantity. 

The  chronic  type  may  succeed  the  acute,  or  appear  like  diarrhoea, 
the  motions  being  faeculent  and  containing  mucus.  Between  exacerba- 
tions, constipation  may  occur.  The  number  of  motions  may  only  be 
twelve  to  fourteen  per  diem.  Gangrenous  complications  may  occur 
at  any  time,  and  chronic  dysentery  may  persist  for  many  years. 

The  latent  type  is  important,  as  the  patients,  though  free  from 
dysenteric  symptoms,  harbour  amoebae  and  act  as  parasite  carriers. 
The  latent  condition  may  lead  to  acute  attacks  or  to  liver  abscess. 

The  mixed  type  occurs  where  amoebic  and  baciilary  dysentery  are 
combined.  There  is  much  fever,  nausea,  and  vomiting.  The  motions 
are  numerous  and  often  very  offensive. 

Treatment. — The  most  modern  method  of  treatment,  due  to 
Leonard  Rogers,  is  by  emetine.  According  to  Castellani  and 
Chalmers,  it  is  w^ell  to  relieve  griping  and  straining  by  either  a  hypo- 
dermic injection  of    morphia  or  by  small  enemata  of  40  minims  of 

^Philippine  Journ.  Sc,  B,  viii,  p.  253. 


SUPPLEMENT  619 

laudanum  in  1  oz.  of  mucilage  of  starch  or  by  using  ^  gr.  morphia 
or  i  gr.  codeine  suppository.  A  dose  of  castor  oil  (5iv  to  5vi)  with 
or  without  a  few  minims  of  liquor  opii  sedativus  or  a  few  doses  of 
saline  may  be  given  during  the  first  twenty-four  hours.  After  the 
castor  oil  has  acted  or  simultaneously,  emetine  treatment  should  be 
commenced  ;  ^  to  i  gr.  of  emetine  hydrochloride,  dissolved  in  sterile 
normal  salt  solution,  is  injected  hypodermically  three  times  a  day  for 
two  or  three  days. 

If  emetine  cannot  be  obtained,  5  gr.  doses  of  ipecacuanha  every 
three  to  six  hours  in  the  form  of  membroids,  or  as  pills  coated  with 
salol  or  keratin,  can  be  substituted. 

After  acute  symptoms  have  disappeared,  intestinal  irrigations  once 
or  twice  daily,  on  alternate  days,  are  useful.  A  solution  of  tannic  acid 
(3  ^o  5  P^^  1,000)  or  of  quinine  bihydrochloride  varying  in  strength 
from  I  in  5,000  to  i  in  750  is  very  slowly  injected  in  quantities  of 
^  to  3  pints  by  means  of  a  long,  soft,  rectal  tube. 

For  gangrenous  dysentery  Castellani  and  Chalmers  state  that 
appendicostomy,  with  irrigation  of  the  whole  lower  bowel  with 
quinine  lotion  (i  in  1,000)  or  collargol  (i  in  500),  is  the  only  chance. 

The  use  of  emetine  should  be  continued  in  smaller  doses  after  the 
dysenteric  symptoms  have  ceased,  in  order  to  prevent  relapses  and  as 
a  possible  safeguard  against  the  development  of  a  liver  abscess. 

Recently  (July,  1914),  Dr.  W.  E.  Deeks^  has  given  an  account  of 
his  successful  procedure  in  dealing  with  the  dysenteries  in  the  Ancon 
Hospital,  Panama  Canal  Zone,  of  which  medical  clinic  he  is  the  chief. 
With  regard  to  amoebic  dysentery  he  advocates  :  (i)  Rest,  to  increase 
the  patient's  resistance ;  (2)  a  generous  milk  diet,  which  is  practically 
all  absorbed  before  it  reaches  the  large  bowel  ;  (3)  saline  or  plain 
water  irrigations,  one  to  three  daily ;  (4)  the  administration  of 
bismuth  sub-nitrate  in  heroic  doses  ;  180  gr.  is  given  mechanically 
suspended  in  about  a  tumbler  of  plain  or  effervescent  water  every 
three  hours,  day  and  night  in  severe  cases,  only  lessening  the  amount 
when  improvement  takes  place.  Mechanical  suspension  in  a  large 
quantity  of  water  is  essential.  When  the  stools  begin  to  decrease  in 
number  and  the  tongue  becomes  clean,  the  number  of  doses  is  reduced 
to  three  or  four  daily.  In  very  chronic  cases  one  or  two  doses  daily 
for  a  month  after  convalescence  are  recommended. 

In  exceptional  cases  of  extreme  emaciation  and  exhaustion, 
showing  marked  toxic  symptoms,  surgical  treatment  is  necessary, 
and  at  Ancon  a  wide,  open  caecostomy  is  performed. 

The  treatment  of  dysentery  with  bismuth  sub-nitrate  has  been  in 
use  for  some  years  at  Ancon.  Latterly,  a  combined  treatment  by 
hypodermic   injections    of   emetine  and  bismuth  sub-nitrate   by  the 

'  Annals  Trop.  Med.  and  ParasitoL,  viii,  pp.  321,  353. 

39 


620  THE   ANIMAL   PARASITES   OF   MAN 

mouth  has  been  used,  and  the  authorities  there  consider  that  it  is 
better  to  combine  the  two  drugs  rather  than  use  each  singly.  Emetine 
probably  acts  as  a  direct  poison  to  the  amoebae,  while  the  bismuth 
probably  acts  by  destroying  the  symbiotic  organisms  necessary  for 
their  growth. 

With  regard  to  preventive  measures,  all  drinking  water  should  be 
filtered  and  boiled,  and  uncooked  vegetables  and  salads  avoided. 
Scrupulous  care  w^ith  regard  to  personal  cleanliness,  and  avoidance 
of  touching  the  mouth  or  lips  after  contact  with  dysenteric  patients, 
are  essential.  Isolation  of  parasite  carriers  is  of  great  use  in  combat- 
ing and  controlling  outbreaks  of  amoebic  dysentery.  The  pollution 
of  soil  and  water  must  be  rigorously  prevented. 

Liver  abscess  due  to  amoebae  must  be  localized  by  exploratory 
punctures,  and  then  opened  and  drained.  Intramuscular  injections 
fo  emetine  hydrochloride,  ^  gr.  to  i  gr.  every  day,  will  reduce  the 
temperature  and  afford   relief. 

Oral  endamoebiasis  has  been  recently  investigated  by  Bass  and 
Johns,  Smith  and  Barrett  and  colleagues  (see  pp.  43,  733).  It  responds 
to  treatment  with  emetine,  and  ^  gr.  of  emetine  hydrochloride 
administered  hypodermically  each  day  is  of  service.  Rinsing  the 
mouth  with  a  solution  of  fluid  extract  of  ipecacuanha  is  also  useful. 

Rogers^  (iQiS)  recommends  a  combined  treatment  of  emetine 
and  streptococcal  vaccines  for  pyorrhoea  alveolaris. 

II.— TRYPANOSOMIASES. 

The  human  trypanosomiases  are  those  occurring  in  Africa,  due  to 
Trypanosoma  ganibiense  and  T.  rhodesiense  and  spread  by  Glossinae, 
and  that  due  to  T.  cnizi,  occurring  in  South  America  and  spread 
by  the  Reduviid  bugs,  Triatoma  spp.  These  trypanosomiases  present 
different  clinical  features  and  are  best  dealt  with  separately. 

African  Sleeping  Sickness. 

Sleeping  sickness,  due  to  Trypanosoma  gamhiense  or  varieties 
thereof,  was  first  reported  from  West  Africa  and  is  now  present,  not 
only  along  the  West  Coast  and  in  Nigeria,  but  throughout  the  Congo 
basin  into  Uganda,  north  of  which  it  exists  in  the  Bahr-el-Ghazal 
province  of  the  Sudan.  In  Nyasaland  and  Rhodesia  a  more  virulent  but 
less  widely  distributed  disease  is  produced  by  Trypanosoma  rhodesiense. 

There  is  a  general  similarity  between  the  two  diseases,  and  the 
symptoms  as  described  by  the  leading  authorities  agree  in  the  main. 
The  malady  due  to  T.  rhodesiense  has  been  known  only  since  1910 
and  the  differences  between  the  malady  due  to  it  and  to  T.  gamhiense 
will  be  indicated. 

^  Ind.  Med.  Gazette^  April,  1915,  I,  p.  121. 


SUPPLEMENT  621 

The  course  of  the  disease  may  be  roughly  divided  into  three 
stages,  the  incubation,  the  febrile  or  glandular,  and  the  cerebral  stage. 

The  exact  incubation  period  is  not  known  with  certainty  in  man. 
Probably,  in  most  cases,  it  does  not  exceed  two  to  three  weeks,  but 
disease  signs  may  not  appear  for  months.  The  bite  of  the  Glossina 
gives  rise  to  local  irritation,  which  may  be  overlooked.  The  irrita- 
tion usually  subsides  in  the  course  of  a  few  days. 

The  febrile,  or  glandular  stage,  is  marked  by  attacks  of  fever  of  an 
intermittent  type.  An  erythematous  eruption  is  often  found  on  Euro- 
peans. This  rash  begins  as  irregularly  shaped  pinkish  patches  which 
clear  in  the  centre  until  a  ring  is  produced.  It  may  occur  on  any 
part  of  the  body  but  is  more  frequent  on  the  trunk.  A  typical 
symptom  is  the  enlargement  of  one  or  more  of  the  lymphatic  glands, 
especially  those  of  the  neck.  A  general,  deep  hyperaesthesia,  known 
as  Kerandel's  sign,  may  be  present,  and  if  the  patient  strikes  a  limb 
against  any  hard  object,  a  feeling  of  acute  pain  is  felt,  the  sensation 
being  slightly  delayed.  As  repeated  attacks  of  fever  increase,  the 
patient  may  become  anaemic.  The  febrile  stage  may  last  for  years, 
and  cure  may  be  brought  about  at  tliis  phase,  but  frequently,  after  the 
febrile  stage  has  lasted  some  time,  the  cerebral  stage  is  reached. 
Tachycardia  is  also  a  symptom.  Auto-agglutination  of  the  red  blood 
corpuscles  is  another  useful  characteristic,  as  it  is  said  to  occur  rarely 
in  other  tropical  diseases,  but  some  workers  doubt  its  value. 

The  cerebral,  or  true  sleeping  sickness  stage  is  marked  by  a  great 
change  in  the  habits  of  the  victim,  who  becomes  apathetic  and  dull, 
careless  and  dirty  in  habits,  and  begins  to  experience  difficulty  in 
walking.  Tremors  of  varying  degrees  of  severity  are  common  and 
the  gait  is  peculiar.  There  is  usually  fever  with  rise  of  temperature 
from  100°  F.  to  104°  F.  in  the  evening,  becoming  subnormal  in  the 
morning.  For  some  days  before  death,  it  often  becomes  permanently 
subnormal.  Congestion  and  oedema  of  the  lungs,  with  patches  of 
pneumonia,  are  not  infrequently  observed  before  death.  The  torpor 
gradually  deepens,  and  the  patient  loses  flesh.  Frequently  the  lips 
swell  and  saliva  dribbles.  The  patient  usually  becomes  comatose  and 
death  ensues.  Mania  and  delusions,  and  psychical  and  physical 
symptoms  resembling  those  found  in  general  paralysis  of  the  insane, 
sometimes  occur,  and  death  may  arise  from  secondary  complications 
such  as  pneumonia  or  dysentery. 

Pathologically,  the  disease  seems*  to  consist  of  a  chronic  inflam- 
mation of  the  lymphatic  system.  The  trypanosomes  reach  the 
lymphatic  glands  which  become  inflamed,  and  gradually  invade  the 
blood  and  the  cerebrospinal  fluid.  Sooner  or  later,  as  a  result  of 
the  lymphatic  disease,  changes  occur  in  the  membranes  and 
substances  of  the  brain  and  spinal  cord.  There  is  round-celled 
perivascular  infiltration  of  the  pia-arachnoid  of  the  brain  and  spinal 


622  THE   ANIMAL   PARASITES   OF   MAN 

cord.  These  changes  cause  compression  of  the  blood-vessels,  and 
so  lessen  the  supply  of  blood  to  the  brain  and  spinal  cord.  Further 
changes  in  the  latter  organs  result  in  the  production  of  the  symptoms 
that  have  given  the  disease  the  name  of  **  sleeping  sickness." 

The  disease  due  to  Trypanosoma  rhodesieiise  generally  runs  a  more 
rapid  course  than  that  due  to  T.  gainbiense.  The  torpor  and  sleepi- 
ness may  not  be  obvious  or  be  very  slight,  and  the  enlargement  of 
the  lymphatic  glands  of  the  neck  also  may  not  be  marked  or  may 
appear  to  be  absent.  The  duration  of  .  the  disease  often  appears 
to  be  from  three  to  six  months. 

Treatment  is  only  of  use  if  commenced  in  the  earlier  stages  of 
the  disease.  The  substances  of  most  value  so  far  are  arsenic  in 
the  form  of  atoxyl  (introduced  by  Wolferstan  Thomas  in  1905)  and 
antimony  in  the  form  of  tartar  emetic.  Castellani  and  Chalmers  and 
Manson  recommend  treatment  by  combining  the  use  of  both  substances. 
The  combined  treatment  is  recommended  not  only  because  both  sub- 
stances have  been  proved  of  service  independently,  but  also  because 
certain  strains  of  trypanosomes  resistant  to  arsenic  are  known,  and 
trypanosomes  can  develop  a  resistance  to  arsenic.  Such  forms,  that 
would  not  be  affected  by  the  atoxyl,  are  left  open  to  attack  by  the 
antimony  salt.  Daniels  also  recommends  combined  arsenic  and 
antimony  treatment,  and  (19 15)  uses  atoxyl  and  antiluetin. 

Atoxyl  is  best  given  intramuscularly  in  10  per  cent,  solution  in  sterile 
normal  saline  solution.     Galyl  is  also  said  to  have  given  good  results. 

Castellani  and  Chalmers  recommend  :  (i)  Hanson's  method  of 
administration  of  atoxyl,  viz.,  2  to  3  gr.  of  atoxyl  are  given  by 
intramuscular  injection  every  third  day  for  at  least  two  years;  or 
(2)  Broden  and  Rodhain's  method,  7^  gr.  of  atoxyl  by  intramuscular 
injection  every  fifth  day.  For  the  combined  therapy  by  atoxyl  and 
antimony  they  recommend  the  following  : — ''  An  atoxyl  injection 
(3  gi'-)  is  given  every  third  day  or  7^  gr.  every  fifth  day,  and  sodio- 
tartrate  of  antimony  (Plimmer's  salt)  is  administered  daily,  2  gr. 
dissolved  in  a  large  quantity,  of  water  (2  pints)  by  the  mouth  or 
by  the  rectum.  Tartar  emetic,  however,  is  best  given  by  intravenous 
injections,  using  solutions  of  i  in  100  or  i  in  1,000.  The  dose  of 
the  drug  to  be  given  is  5  to  10  eg.  per  injection.  It  is  important 
that  none  of  the  fluid  of  the  injection  should  escape  into  the 
surrounding  tissues,  as  a  violent  inflammation  may  result.  These 
injections  should  be  administered  monthly  on  ten  consecutive  days 
for  a  long  period." 

Macfie  and  Gallagher  (1914)  injected  6  gr.  of  atoxyl  intra- 
muscularly every  week  in  cases  infected  with  7 .  nlgeriense  in  the  Eket 
district  of  Southern  Nigeria. 

Large  doses  of  atoxyl  were  often  said  to  cause  distressing  results 


SUPPLEMENT  '  623 

such  as  optic  atrophy,  and  when  the  onset  of  such  occurred  the  drug  was 
usually  discontinued.  However,  Daniels^  (July»  iQ^S)  points  out  that 
eye  troubles,  such  as  iridocyclitis,  are  symptoms  of  trypanosomiasis.  - 

Other  arsenical  preparations  such  as  soamin  and  arsenophenylglycin 
have  been  used,  but  less  successfully  than  atoxyl.  Fowler's  solution, 
well  diluted,  has  been  given  by  the  mouth  when  treatment  by  injection 
was  not  possible,  the  doses  commencing  with  5  minims  and  increasing 
to  15  minims. 

Salvarsan  and  neo-salvarsan  have  also  been  tried  for  sleeping 
sickness.  Plimmer  recommended  powdered  antimony  suspended 
in  sterile  olive  oil.  Ranken  used  precipitated  metallic  antimony  in 
normal  saline  solution  injected  intravenously. 

Laveran  and  Thiroux  have  recommended  a  combined  treatment 
of  atoxyl  and  an  inorganic  salt  of  arsenic  such  as  orpiment.  The 
orpiment  is  given  as  pills,  in  doses  of  2  gr.  of  orpiment  two  or  three 
times  daily.  Opium  is  added  to  the  orpiment  to  prevent  diarrhoea. 
This  treatment  is  said  to  have  been  used  in  man  with  good  results. 

Trypanosoma  rhodesieiise  seems  less  amenable  to  treatment  than 
T .  gambiense. 

The  main  preventive  measures  seem  to  he  in  segregation  of  the 
sick  in  areas  not  infested  with  Glossinae,  and  in  measures  against  these 
flies,  such  as  bush  clearing  and  destruction,  to  some  extent,  of  proved 
reservoirs  in  big  game. 

South  American  Trypanosomiasis. 

The  chief  clinical  features  of  the  trypanosomiasis  occurring  ni 
Brazil  have  already  been  indicated  (see  p.  87).  With  regard  to 
treatment,  according  to  Castellani  and  Chalmers  the  indications  are 
the  same  as  those  for  African  trypanosomiasis,  together  with  treat- 
ment for  hypothyroidism.  Preventive  measures  are  directed  against 
the  Reduviid  bug,  Triatoma  rnegista,  that  transmits  the  disease.  The 
bugs  occur  in  numbers  in  the  cracks  of  the  houses  of  the  poor  of 
Minas  Geraes,  and  may  be  destroyed  by  sulphur  fumigation,  lime- 
washing  or  whitewashing. 

III.-FLAGELLATE  DIARRH(EA  AND  DYSENTERY. 

The  chief  causal  agents  are  Trichomonas  hominis  {T.  intestinal  is), 
CJi ilomastix  (Tetramitns)  mesnili  and  allied  organisms  (see  pp.  54  to  57), 
and  Lamblia  intestinalis  (see  pp.  57  to  60  and  Appendix  pp.  734  to  736). 

These  parasites  and  the  associated  diarrhoeas  occur  in  temperate 
as  well  as  in  warm  climates.  Probably  some  of  the  diarrh(jeas  in  India 
are  thus  caused.  The  same,  or  similar  parasites  occur  in  various 
Muridae,  especially  rats  and  mice,  which  may  act  as  reservoirs. 

'  lourn.  Trop.  Med.  and  Hyg.y  xviii,    p.   157. 


624  THE   ANIMAL   PARASITKS   OF   MAN 

(i)  Mello-Leitao^  (1913);  writing  from  Rio  de  Janeiro,  states  that 
there  is  a  primary  flagellate  dysentery,  due  to  Trichoinonas  iniestinalis 
(Leiickart)  and  to  Lamblia  iniestinalis  (Lambl),  either  separately  or  in 
combination.  He  considers  it  a  benign  disease,  and  the  most  frequent 
form  of  dysentery  in  young  children.  Trichomonas  and  Lamblia 
were  found  to  be  pathogenic  to  children  under  3  years  of  age. 

EscomeF  (191 3)  collected  152  cases  of  dysentery  in  Peru  due 
solely  to  Trichomonas.  Examination  of  the  reservoirs  containing 
the  water  used  for  drinking  purposes  showed  the  presence  of  Tricho- 
monas. After  the  reservoirs  were  cleaned  no  more  Trichomonas  was 
found  and  the  cases  of  dysentery  ceased. 

Brumpt^  (191 2)  described  a  colitis  due  to  Trichomonas  intestiiialis 
in  a  patient  returned  from  Tonkin. 

Cases  of  infection  by  Cliiloniastix  (Tetramitus)  niesnili,  with  colitis 
or  dysenteric  symptoms,  are  recorded  by  Brumpt  (1912)  from  France, 
and  by  Nattan-Larrier^  (191 2)  from  the  Ivory  Coast  respectively. 

Marques  da  Cunha  and  Torres'^  (1914)  describe  five  cases  of 
chronic  diarrhoea  in  Brazilian  children  due  to  the  Chilomastix 
(Tetramitus). 

GabeP  (1914)  described  a  case  of  seasonal  diarrhcjea  contracted  in 
Tunis  and  caused  by  a  Tetramitid  parasite  which  he  named  Difdmus 
tnnensis,  as  the  discoverer  considered  that  it  lacked  an  undulating 
membrane  in  its  large  cytostome. 

Derrieu  and  Raynaud^  (i9H)  I'ecord  a  case  of  chronic  dysentery 
in  Algeria  due  to  a  Trichomonad  possessing  an  undulating  membrane 
and  five  free  flagella.  The  parasite  was  named  Hexamastix  ardin- 
delteilif  but  the  generic  name  Hexamastix  is  pre-occupied.  Chatterjee's 
Pentatrichomonas  bengalensis  (191 5)  is  possibly  the  same  organism. 

Treatment. — Escomel  (1913),  finding  ipecacuanha  and  calomel 
useless,  recommends  turpentine  for  Trichomonad  dysentery.  Two  to 
4  grm.  of  essence  of  turpentine  in  an  emulsion  are  given  by  the  mouth, 
and  enemata  containing  15  to  20  drops  of  turpentine  emulsified  in 
the  yolk  of  an  egg  to  which  is  added  a  little  water  and  tincture  of 
opium.  Derrieu  and  Raynaud  found  this  treatment  effective  in  Algeria. 
Smithies^  (19 12)  reports  two  cures  of  cases  of  severe  dyspepsia,  in 
which  Trichomonads  were  found  in  the  stomach  contents,  after 
administration  of  a  single  dose  of  50  to  60  gr.  of  thymol,  given  at 
bed-time,  together  with  2  gr.  of  calomel,  and  followed  by  an  ounce 
of    Carlsbad   salts    in    the    morning.     The    patients   came    from    the 

'  BriL  /ourn.  Child}  en's  Diseases,  x,  p.  60. 
-  Bttll.  Soc.  Path.  Exol.,  vi,  p.  120.  '  Ibid.,  v,  p.  725. 

*  Ibid.,  V,  p.  495.  5  Btazil  Medico,  xxviii,  p.  269. 

^  Arch.f.  Protistenkimde,  xxxiv,  p.  i.  ^  Bull.  Soc.  Path.  Exot.,  vii,  p.  571. 

^  Amer.  fottrn.  Med.  Sci.,  cxliv,  p.  82. 


SUPPLEMENT  625 

Southern  United  States,  and  had  been  in  the  habit  of  drinking 
unfiltered  surface  water  in  the  locaHties  in  which  they  Hved.  Mello- 
Leitao^  used  magnesium  sulphate  and  water  or  milk  diet.  Sometimes 
enemata  of  coUargol  (i  per  cent.)  or  electrargol  were  required. 
Rosenfeld  recommended  calomel.  Methylene  blue  has  also  been 
tried.  Recently,  EscomeP  (1914)  recommends  enemata  of  an  aqueous 
solution  of  iodine  (i  per  1,000)  and  farinaceous  diet.  Lynch^  (19 15), 
working  in  South  Carolina,  recommends  a  mouth  wash  of  saturated 
solution  of  bicarbonate  of  soda  three  times  daily  in  oral  infections. 
A  similar  solution  was  used  as  a  douche  in  vaginal  trichomoniasis. 

Stiles  (19 1 3)  points  out  that  when  amoebae  or  flagellates  are  found 
in  a  large  percentage  (10  to  40,  or  even  60)  of  the  members  of  a 
community,  means  should  be  taken  to  improve  the  methods  for  the 
disposal  of  the  dejecta,  so  that  the  food-supply  may  be  carefully 
protected  against  faecal  contamination.  Cysts  of  the  parasites  may  be 
air-borne  or  conveyed  to  food  on  the  bodies  of  house-flies. 

(ii)  Lainhlia  intestinalis  in  man  may  cause  diarrhoea  with  dysenteri- 
form  stools.  The  diarrhoea  may  be  of  a  chronic  recurrent  character. 
The  flagellate,  or  a  variety  of  it,  is  fairly  common  in  the  digestive  tract 
of  rats  and  mice. 

Mathis^  (1914)  gives  an  interesting  account  of  cases  in  Tonkin. 
In  a  child,  aged  3,  the  stools  were  at  first  glairy  and  blood-stained, 
containing  many  encysted  Lamblia.  The  child's  home  was  infested 
with  mice.  In  another  case,  the  house  of  the  patient  harboured 
numerous  rats. 

According  to  Mathis,  prognosis  is  favourable,  but  emetine  hydro- 
chloride is  without  action  on  Lamblia.  Prowazek  and  Werner^ 
(1914),  however,  state  that  emetine  will  act  upon  the  flagellates,  but 
not  upon  the  cysts.  They  recommend  uzara  (two  tablets,  three  times 
daily)  and  extract  of  male  fern  as  useful  in  certain  cases.  Martin 
Mayer  (1914)  found  emetine  hydrochloride  successful  in  a  case  in  the 
Hamburg  Seamen's  Hospital,  but  Assmy  (19 14)  points  out  that  a 
suitable  diet  and  daily  doses  of  magnesium  sulphate  are  sufficient, 
in  his  experience,  to  effect  an  improvement,  and  he  doubts  the  specific 
action  of  emetine.  Escomel  (1914)  recommends  milk  diet,  then 
calomel  succeeded  by  castor  oil. 

According  to  Noc,  Lamblia  may  also  be  water-borne.  Healthy 
carriers  of  Lamblia  cysts  are  known.  Food  should  be  protected 
from  being  soiled  by  rats  and  mice. 


'  Brit.  Journ.  Children's  Diseases,  x,  p.  60.  ^  ^^^//^  Soc.  Path.  Exot.,  vii,  p.  657. 

3  Atner.  Journ.  Trop.  Dis.  and  Prevent.  Med.,  ii,  p.  627. 

*  Bull.  Soc.  Med.  Chirurg.  Indo-Chine,  v,  p.  55. 

5  Beihefte  z.  Arch.  f.  Schiffs-  ti.  Tropen-Hyg.,  xviii,  5,  p.  155. 


626  THE   ANIMAL    PARASITES   OF    MAN 

lY.— LEISHMANIASES. 

A.     Kala-azar. 

(i)   ^^  Indian  "  Kala-azar  cine  to  Leishmania  donovani. 

Indian  kala-azar  due  to  Leishmania  donovani  is  a  very  fatal  disease 
with  a  rate  of  mortality  varying  from  70  to  98  per  cent,  of  the  cases. 

The  incubation  period  is  very  variable  and  the  early  symptoms 
not  well  defined.  The  incubation  period  seems  to  range  from  three 
weeks  to  several  months  after  exposure  to  infection.  The  onset  seems 
to  commence  with  a  rigor  and  attack  of  irregular,  remittent  fever,  which 
may  show  two  remissions  per  day  in  a  four-hourly  temperature  chart. 
Rogers  considers  the  daily  double  remission  almost  diagnostic.  The 
duration  of  this  first  attack  is  from  two  to  six  weeks.  The  spleen 
and  liver  enlarge,  especially  the  former,  and  are  painful  and  tender. 
Towards  the  end  of  the  time  the  temperature  declines  and  the  first 
period  of  the  disease  ends.  After  this  period  an  apyrexial  interval 
occurs,  which,  after  some  weeks,  ends  in  an  attack  of  fever  resembling 
the  first.  Periods  of  pyrexia  and  apyrexia  alternate.  Anaemia  com- 
mences and  asthenia  appears  and  deepens  steadily.  The  patient  is 
now  thin  and  wasted,  the  abdomen  much  swollen  and  protuberant, 
the  ribs  show  clearly,  the  limbs  are  wasted  and  skin  and  tongue 
darker  than  normal.  In  Europeans  the  skin  is  of  a  remarkable  earthy 
hue,  and  in  natives  of  India  darker  than  normal,  approaching  black. 
Intestinal  disturbances,  often  in  the  form  of  very  obstinate  and 
intractable  diarrhoea  or  dysenteric  attacks,  are  common.  Papular 
eruptions  often  appear,  particularly  on  the  thighs  ;  haemorrhages  also 
may  occur.  The  disease  lasts  for  periods  varying  from  seven  months 
to  two  years,  and  usually  ends  fatally. 

Treatment,  unfortunately,  has  not  been  very  successful  up  to  1915. 
Manson  has  reported  two  cases  of  cure  by  intramuscular  injections  of 
atoxyl  daily  or  every  other  day  in  doses  of  3  gr.  Rogers  has  advocated 
large  doses  of  quinine,  60  to  90  gr.  daily  until  the  temperature  falls  and 
then  20  gr.  daily.  Castellani  and  Chalmers  consider  the  best  results 
are  obtained  by  large  doses  of  quinine  given  intramuscularly,  supple- 
mented by  a  course  of  quinine  cacodylate  injections  or  atoxyl  injec- 
tions. Tartar  emetic  should  be  tried  (see  pp.  627,  629),  especially 
as  L.  Rogers  (July,  191 5)  has  had  promising  results  in  ten  cases. 
Castellani  (1914)  and  Mackie  (1915),  have  also  had  successful  results. 
Leishman  states  that  the  administration  of  red  bone-marrow,  either 
raw  or  in  the  form  of  tablets,  may  be  beneficial.  Good  nursing  and 
careful  diet  are  essential,  and  diarrhoea  or  dysentery  must  receive  the 
appropriate  treatment. 

With  regard  to  preventive   measures,  the   extermination  of   bugs 


SUPPLEMENT  627 

and  other  biting  insects  seems  to  be  of  most  service.  Domestic  and 
personal  cleanliness  is  of  great  importance.  Patients  should  be 
segregated.  It  would  probably  be  as  well  if  houses  in  which  many 
cases  of  kala-azar  occurred  were  destroyed.  Dodds  Price,  in  Assam 
tea  gardens,  moves  the  coolie  lines  300  to  800  yards  from  old  infected 
ones,  with  satisfactory  results. 

(ii)     Infantile  Kala-azar  due  to  Leishmania  infantum. 

This  malady  is  found  among  children,  rarely  in  adults,  along  the 
Mediterranean  littoral. 

The  disease  commences  insidiously  and  is  often  unrecognized  until 
some  intestinal  disturbance  occurs.  The  spleen  is  then  found  to  be 
somewhat  enlarged,  and  the  case  has  often  been  regarded  as  one  of 
malaria.  The  child  becomes  anaemic,  suffers  from  diarrhoea,  alternat- 
ing with  constipation,  and  has  attacks  of  irregular  fever.  The  spleen 
continues  to  enlarge  and  protrudes  from  under  the  cover  of  the  ribs. 
Haemorrhages  from  the  nose  and  gums  and  into  the  skin  occur. 
Anaemia  and  wasting  set  in.  The  abdomen  then  becomes  very 
enlarged.  The  child  becomes  much  less  active  both  physically 
and  mentally,  and  looks  prematurely  old.  Death  often  occurs  from 
exhaustion,  though  some  cases  of  spontaneous  recovery  are  known. 

Treatment  up  till  recently  has  been  unsatisfactory.  Some  of  the 
remedies  tried,  as  quoted  by  Castellani  and  Chalmers,  are  15  eg.  doses 
of  atoxyl,  benzoate  of  mercury  (2  to  4  mg.  as  a  daily  injection), 
thiarsol  (5  to  15  mg.  by  subcutaneous  injection),  salvarsan,  etc. 
Recently  Cristina  and  Caronia  (1915)^  have  given  repeated  intravenous 
injections  of  i  per  cent,  aqueous  solution  of  tartar  emetic,  the  dose 
varying  from  2  to  10  eg.  The  treatment  in  various  cases  has  lasted 
from  15  to  40  days. 

Prophylactic  measures  seem  to  lie  in  the  destruction  of  infected 
dogs  and  diminishing  the  breeding  of  fleas  (see  p.  in). 

B.  Oriental  Sore,  due  to  Leishmania  tropica. 

Oriental  sore,  known  under  many  other  names  (see  p.  107),  is  a 
local  infection  of  the  skin  due  io  Leishmania  tropica.  The  incubation 
period  varies  from  a  few  days  to  some  weeks,  or  even  months,  and 
then  one  or  several  small  itching  papules  appear.  Each  spot  becomes 
red  and  shotty,  the  papules  increase  slowly  in  size  and  the  surface 
becomes  covered  with  papery  scales.  After  a  variable  time,  usually 
not  exceeding  three  to  four  months,  ulceration  occurs  and  a  yellowish 
secretion  is  exuded  that  soon  dries  into  a  scab.  Under  the  scab 
ulceration  continues  by  erosion  of  the  edges,  and  subsidiary  sores 
arise  around  the  parent  ulcer  and  usually  fuse  with  it.  Healing  com- 
mences after  six  to  twelve  months.     Granulation  begins  at  the  centre 

lull.  Soc.  Path.  Exot.y  viii,  p.  63. 


628  THE   ANIMAL   PARASITES   OF   MAN 

and  spreads  outwards,  and  when  healing  is  complete,  a  depressed, 
whitish  or  pinkish  scar  remains. 

Many  treatments  for  Oriental  sore  have  been  devised  but  do  not 
seem  particularly  satisfactory.  Castellani  and  Chalmers  state  that 
the  scabs  should  be  removed  by  boracic  acid  fomentations,  and  the 
ulcers  thoroughly  disinfected  once  or  twice  daily  with  a  i  per  i,ooo 
solution  of  perchloride  of  mercury,  after  which  an  ordinary  antiseptic 
ointment  is  applied. 

The  use  of  permanganate  of  potash  has  been  advocated  both  by 
French  and  English  doctors.  Both  large  and  small  sores  can  be 
treated.  The  patient's  skin  around  the  sore  is  protected  by  a  thick 
layer  of  vaseline,  and  the  surface  of  the  ulcer  powdered  with  potassium 
permanganate,  which  is  kept  in  position  by  a  pad  of  gauze  and  a 
bandage.  The  treatment  is  said  to  cause  great  pain  for  six  to  eight 
hours,  but  at  the  most,  three  treatments  are  necessary  before  the  sore 
becomes  a  simple  ulcer,  well  on  the  way  to  healing.  The  perman- 
ganate may  also  be  used  in  ointment.  Excision  of  the  ulcer  when 
small  is  advisable  when  the  site  of  the  ulcer  permits  of  this.  According 
to  Manson,  reports  on  treatment  by  radium,  salvarsan  and  carbon 
dioxide  snow  are  decidedly  promising.  Mitchell  (1914)^  reports 
favourably  on  the  use  of  carbon  dioxide  snow  in  the  form  of  a  pencil, 
in  India.  In  Brazil  several  workers  (1914)  record  successful  results 
from  the  intravenous  injection  of  a  i  per  cent,  solution  of  tartar 
emetic  in  distilled  water.  Low  (191 5)  has  successfully  treated  a  case  by 
•direct  local  application  of  tartar  emetic.  Row  (191 2)  has  treated  cases 
of  Oriental  sore  by  inoculation  of  killed  cultures  of  the  causal  organism. 

As  the  disease  is  very  contagious,  the  slightest  wound,  and  any 
insect  bite,  should  be  thoroughly  disinfected  with  5  per  cent,  carbolic 
acid  or  iodine.  Destruction  of  bugs,  lice,  and  other  biting  insects 
should  be  enforced.  As  dogs  may  contract  the  disease  (see  p.  108), 
it  is  well  not  to  allow  them  in  the  house  and  not  to  encourage  undue 
contact  with  them. 

Naso-oral  Leishmaniasis  {Espundia)  due  to  Leishmania  tropica. 

This  form  of  Leishmaniasis  has  been  reported  from  South  America 
and  recently  by  Christopherson^  {1914)  fi'oi^i  the  Sudan.  In  South 
America  it  is  often  called  Espundia,  also  Buba  and  Forestal  Leish- 
maniasis. The  primary  lesion  is  found  usually  on  the  forearms, 
legs,  chest  or  trunk.  This  ulcer  is  of  the  Oriental  sore  type,  and 
after  some  months,  or  even  as  long  as  two  years,  heals  up,  leaving  a 
thick  scar.  While  the  ulcer  is  open,  or  more  often  after  it  has  healed, 
lesions  appear  on  the  mucosa  of  the  mouth  and  nose.     The  hard  and 

'  Journ.  Roy.  Army  Med.  Corps,  xxiii,  pp.  440-446  (see  Trap.  Dis.  BiilL^v,  No.  5,  p.  276). 
"^  Annals  Tiop.  Med.  and  Paras itoL,  viii,  p.  485. 


SUPPLEMENT  629 

soft  palate,  gums  and  lips  all  may  be  attacked.  The  mucosa  of  the 
nose  is  usually  attacked  and  the  cartilages  become  destroyed,  produc- 
ing great  deformity.  In  bad  cases  the  pharynx  and  larynx  may 
become  infected. 

Till  recently  it  was  believed  that  treatment  was  of  little  use  unless 
the  case  could  be  investigated  early.  Escomel  considered  that  if  the 
primary  cutaneous  lesion  w^as  excised  or  destroyed,  further  progress 
of  the  disease  was  prevented.  When  lesions  have  appeared  on  the 
mucosa  of  the  mouth  or  nose,  little  could  be  done.  The  ulcers  might 
be  cauterized  and  mild  antiseptic  mouth  washes  used. 

In  1913  Vianna,  working  in  Brazil,  introduced  treatment  by  tartar 
emetic,  which  is  now  becoming  more  wadely  known  and  proving 
efficacious.  Carini^  (1914)  applies  it  thus.  Tartar  emetic  (that  is, 
potassium  antimonyl  tartrate)  in  i  per  cent,  aqueous  solution  is  intro- 
duced slowly  into  a  vein,  such  as  the  vein  at  the  bend  of  the  elbow, 
in  doses  of  5  to  10  c.c.  daily  or  on  alternate  days  according  to  the 
tolerance  of  the  patient  to  the  drug.  Eighteen  to  forty  injections 
have  been  used.  In  some  of  the  memoirs  on  the  subject,  the  drug  is 
referred  to  as  antimony  tartrate. 

The  course  of  the  disease  is  chronic  and  may  last  for  twenty  to 
thirty  years,  death  usually  resulting  from  some  intercurrent  disease. 

At  present  the  actual  transmitter  of  Espundia  is  not  known  with 
certainty.  Various  sand-flies  (Simulidae)  have  been  suspected  of 
transmitting  the  disease,  though  so  far  proof  is  wanting.  It  has  also 
been  suggested  that  the  natural  food  sources  of  some  Simulidae  known 
to  bite  man,  namely,  certain  snakes^  and  lizards,^  are  possible 
reservoirs  of  the  disease. 

Prophylactic  measures  would  seem  to  consist  in  the  immediate 
disinfection  of  insect  bites  by  tincture  of  iodine,  and  by  avoidance  of 
areas  known  to  be  infested  with  snakes  and  lizards,  and  insects  that 
prey  on  them  and  man  indifferently.  The  destruction  of  the  primary 
lesion  as  soon  as  detected  is  essential,  and  the  isolation  of  advanced 
cases  of  the  disease  seems  advisable. 

Y.— SPIROCHiETOSES. 

A.    Relapsing     Fevers. 

The  relapsing  fevers  of  Europe  and  of  America,  due  to  Spirochceta 
recur  reiitis  and  S.  novyi  (probably  a  race  of  S.  recurrent  is),  present 
much  the  same  symptoms,  w^hich  differ  in  some  respects  from  those 

•  Bu//.  Soc.  Path.  Exot.,  vii,  p.  277. 

2  Lindsay  (1914),  Trans.  Soc.  Trop.  Med.  and  Hyg.,  vii,  p.  259. 

'  Sergent  (Ed.  and    Et.),   Lemaire   and  Senevet   (1914),    Btdl.    Soc.    Path.   Exot.y    vii, 
P-  577. 


630  THE   ANIMAL  PARASITES   OF   MAN 

due  to  S.  duitoni,  the  excitant  of  ''  tick  "  or  ''  relapsing  "  fever  in 
Africa  (see  pp.  1 16-122). 

The  incubation  period  of  S.  recnrrentis  varies  from  two  to 
twelve  days,  during  which  time  a  very  slight  indisposition  may  be 
noticed.  The  onset  is  usually  sudden,  with  severe  headache,  pains 
in  the  back,  limbs  and  stomach  and  a  feeling  of  weakness.  There 
is  a  rise  of  temperature  to  103°  F.  or  104°  F.,  and  the  temperature 
continues  high  till  about  the  sixth  or  seventh  day.  The  skin  is 
yellowish,  hot  and  damp  ;  a  rash,  disappearing  on  pressure,  may  occur 
on  the  trunk  and  legs,  nausea  is  always  present  and  thirst  is  usual. 
The  liver  and  spleen  both  enlarge.  The  number  of  respirations  and 
pulse-rate  become  increased.  On  the  sixth  or  seventh  day  a  crisis 
occurs.  There  is  violent  perspiration,  with  a  rapid  fall  of  tempera- 
ture, pulse  and  respiration  become  normal  and  the  patient  sleeps  and 
awakes  better.  Improvement  continues  for  some  days,  and  recovery 
may  ensue,  but  usually  about  the  fourteenth  day  relapse  occurs, 
lasting  usually  three  or  four  days.  A  second  relapse  is  unusual. 
Numerous  complications  are  known,  e.g.,  bronchitis,  pneumonia, 
diarrhoea  and  dysentery. 

With  regard  to  treatment,  the  specific  appears  to  be  salvarsan. 
Castellani  and  Chalmers  recommend  salvarsan  administered  intra- 
venously. Intramuscular  inoculations  (for  example,  into  the  buttock) 
of  a  suspension  of  "  606  "  in  oil  can  also  be  given.  The  drug  is  very 
efficacious,  but  large  doses  should  not  be  given.  An  intravenous 
injection  of  4  or  5  gr.  does  not  give  rise  to  unpleasant  symptoms 
but  is  sufficient  to  effect  a  cure. 

The  incubation  period  for  the  American  form  of  the  disease  is  at 
least  five  to  seven  days,  and  the  first  attack  lasts  about  five  to  six  days. 
The  treatment  is  by  salvarsan  as  detailed  previously. 

As  relapsing  fever  is  spread  by  body  lice  and  possibly  by  bugs, 
preventive  measures  are  directed  against  these  insects.  Strict 
cleanliness  of  person,  clothing,  bedding  and  dwellings  is  essential. 
Furniture,  e.g.,  wooden  bedsteads,  liable  to  harbour  such  insects 
should  not  be  used. 

The  principal  and  best-known  relapsing  fever  of  Africa  is  that 
excited  by  Spirochcvta  dutioni,  and  transmitted  to  man  by  ticks,  chiefly 
Ornithodorus  monbata.  The  incubation  period  is  usually  about 
seven  days  but  may  be  longer.  The  patient  is  dull  and  lethargic, 
perspires  freely  and  is  often  constipated.  The  temperature  rises  to 
103°  F.  or  105°  F.,  there  is  headache,  pains  in  the  back  and  limbs, 
general  chilliness  and  great  pain  in  the  region  of  the  spleen,  which 
often  enlarges.  The  symptoms  become  w^orse,  there  is  a  fall  of 
temperature  with  improvement  in  the  morning,  and  a  rise,  with 
increase    of    pain,  in  the  evening.     Spirochaetes  are    now  found  in 


SUPPLEMENT  63 1 

the  blood  in  greater  numbers.  The  symptoms  last  three  to  four  days 
and  end  in  a  crisis  with  profuse  sweating  and  fall  of  temperature 
below  normal.  The  day  before  the  crisis  there  is  a  pseudo-crisis, 
when  the  temperature  falls  but  there  is  no  improvement.  The 
patient  is  left  weak  and  tired.  Recovery  may  follow,  but  more  usually 
a  relapse  occurs.  The  intermission  period  varies  ;  five  to  eight  days 
is  common.  The  symptoms  of  the  relapses  are  like  those  of  the  first 
attack.     The  number  of  relapses  varies,  five  to  eleven  may  occur. 

The  treatment  recommended  is  by  salvarsan,  as  for  the  European 
relapsing  fever. 

With  regard  to  prophylaxis,  localities  w^here  ticks  abound  must 
be  avoided  and  the  parasites  themselves  destroyed.  Native  huts 
should  be  avoided.  Mosquito  nets,  a  bed  well  off  the  ground  and 
the  use  of  night  lights  are  advised  by  Manson  to  avoid  attacks  by 
ticks,  which  are  often  nocturnal  in  their  habits. 

In  North  Africa  (Algeria,  Tunis,  Tripoli,  Egypt),  and  sometimes 
in  the  Anglo-Egyptian  Sudan,  a  spirochaetosis  due  to  6>.  berhera 
occurs.  According  to  Castellani  and  Chalmers,  the  incubation  period 
v^aries  somewhat.  The  fever  reaches  its  height  during  the  first  twenty- 
four  hours,  and  afterwards  shows  a  morning  remission.  Jaundice 
is  often  absent,  but  there  may  be  hepatic  tenderness  and  splenic 
enlargement.  One  or  two  relapses  usually  occur.  The  treatment  is 
on  the  same  lines  as  for  the  other  spirochaetal  fevers.  Sergent  and 
Gillot^  (1911)5  working  at  the  Institut  Pasteur  of  Algeria,  have  had 
good  results  by  using  injections  of  salvarsan  in  doses  of  075  to 
T"o  eg.  per  kilogramme  weight  of  the  patient.  The  prophylactic 
measures  are  directed  against  lice  and  other  biting  insects.  Personal 
cleanliness  is  most  necessary. 

In  Asia,  a  relapsing  fever,  due  to  the  spirochaete  named  S.  carteri 
by  Manson  in  1907,  producing  a  mortality  of  about  18  per  cent., 
occurs.  The  symptoms  have  a  general  resemblance  to  those  produced 
by  S.  recurreiitls,  but  on  the  fall  of  temperature  to  subnormal  on  the 
sixth  or  seventh  day,  when  profuse  perspiration  and  polyuria  occur, 
instead  of  improvement  following,  the  patient  often  becomes  collapsed, 
with  a  clammy  skin  and  feeble  pulse.  Improvement  is  slow.  The 
first  relapse  occurs  about  the  fourteenth  day  of  the  attack,  when  the 
temperature  may  be  higher  than  for  the  first  attack.  There  are 
seldom  more  than  four  relapses.  The  treatment  is  by  salvarsan,  of 
which  doses  of  not  more  than  5  gr.  intravenously  should  be  given. 
Sudden  heart  failure  being  common,  Castellani  and  Chalmers  state 
that  cardiac  stimulants  should  be  given.  Prophylaxis  is  the  same  as 
for  European  relapsing  fever. 

*  Bu//.  Soc.  Path.  Exoi.,  iv,  p.  440. 


632  THE   ANIMAL   PARASITES   OF   MAN 

B.  Yaws  or  Framboesia  tropica. 

Yaws  is  essentially  a  tropical  disease,  though  it  is  found  in  the 
tropical  and  subtropical  zones  in  all  parts  of  the  world,  except  in  the 
mountains  and  cold  districts.  In  1905,  Castellani  found  the  causal 
organism,  Treponema  pertenue  (sometimes  called  Spirochccta  pertennis) 
(see  p.  127).  The  disease  shows  three  periods  :  (i)  The  primary  stage, 
consisting  of  the  development  of  the  primary  lesion  or  papule,  which 
is  usually  extragenital.  The  papule  dries  into  a  crust  beneath  which 
an  ulcer  lies.  (2)  The  secondar}^  or  granulomatous  stage,  which  com- 
mences from  one  to  three  months  after  the  primary  lesion  is  first  seen. 
It  consists  of  a  general  eruption  of  small  papules,  some  of  which  enlarge 
and  become  granulomatous  nodules  covered  with  a  yellowish  crust. 
They  are  common  on  the  limbs  and  face.  (3)  The  tertiary  stage,  in 
which  deep  ulcerations  and  gummatous  nodules  appear.  Any  of  the 
tissues  may  be  involved.  Osseous  lesions  may  occur.  The  disease 
does  not  appear  to  be  hereditary  ;  it  is  usually  spread  by  contact. 

The  best  treatment  appears  to  be  by  salvarsan  or  neo-salvarsan. 
Castellani  and  Chalmers  recommend  intramuscular  and  intravenous 
injections.  For  intramuscular  injection  an  alkaline  or  neutral  solu- 
tion of  the  drug  is  preferable,  or  a  suspension  of  the  drug  in  oil 
may  be  used.  The  dose  varies  from  o'3  to  0*5  grm.,  according  to 
the  age  and  sex  of  the  patient.  For  use  intravenously,  a  slightly 
smaller  dose  is  required.     Galyl  is  also  being  used. 

In  countries  where  frambcesia  is  endemic,  slight  skin  abrasions 
should  be  carefully  treated  with  antiseptics.  Yaws  patients  should  be 
isolated  till  cured,  and  their  dwellings  and  personal  possessions 
disinfected. 

C.     Syphilis. 

Syphilis,  due  to  Treponema  pallidum  (sometimes  called  Spiroduvta 
pallida),  is  prevalent  throughout  the  tropics  as  well  as  in  temperate 
zones.  The  disease  is  amenable  to  treatment  by  salvarsan  and  neo- 
salvarsan,  for  administration  of  which  see  relapsing  fever  and  yaw^s. 
Galyl  is  also  being  used  with  favourable  results.  Lambkin's  mercury 
cream  has  been  found  useful  in  treating  numerous  cases  in  Uganda. 
The  life-history  of  the  parasite  is  given  on  p.  124,  and  further  medical 
details  hardly  come  within  the  purview  of  this  book. 

D.     Bronchial  Spirochaetosis. 

Bronchial  spirochaetosis,  due  to  Spirochceta  hronchialis  (see 
pp.  122,  739)  is  probably  of  wide  distribution  in  the  tropics.  The 
spirochaetes  have  been  found  in  cases  of  chest  complaints,  especially 
those  with  bronchitic  symptoms.  The  disease  may  be  suspected  in 
atypical  cases  of  pneumonia  and  bronchitis,  and  may  be  mistaken  for 
incipient  phthisis. 


SUPPLEMENT  633 

Chalmers  and  O'Farrell^  (1913);  writing  from  Khartoum,  re- 
commended rest  in  bed,  good  food  and  ventilation,  coupled  with 
treatment  by  arsenic  in  some  form,  preferably  associated  with  glycero- 
phosphates. These  may  be  given  by  the  mouth,  or  intramuscularly 
as  an  injection  of : — 

Sodium  cinnamate  ...  ...  ...  ...       0*05  grm. 

Sodium  cacodylate  ...  ...  ...       o'lo     ,,' 

Sodiutn  glycerophosphate      ...  ...  ...       o'lo      ,, 

Taylor^  (1913-14),  writing  from  Entebbe,  Uganda,  prescribes 
arsenious  acid  by  the  mouth  in  increasing  doses.  Creosote  has  been 
used  in  West  Africa. 

YL— MALARIA. 

Malaria,  known  also  under  the  names  of  ague,  paludism,  marsh 
fever,  remittent  fever,  intermittent  fever  and  climatic  fever,  among 
others,  is  a  very  widely  spread  disease.  It  is  most  prevalent  in  the 
equatorial  regions  and  gradually  diminishes  north  and  south  of  the 
equator.  The  various  malarial  parasites  (see  pp.  155  to  172)  are  spread 
by  species  of  Anophelines,  and  hence  malaria  is  present  in  districts 
favourable  to  these  intermediate  hosts,  that  is,  in  places  where  there 
is  a  considerable  amount  of  atmospheric  moisture  and  rain,  as  well 
as  heat. 

The  principal  malarial  parasites  are  :  Plasmodium  vivax,  the  agent 
of  simple  tertian  fever ;  Plasmodium  uialarice,  the  parasite  of  quartan 
malaria,  and  Laverania  malarice  or  Plasmodium  falciparum,  producing 
malignant  tertian  or  sub-tertian  malaria  (and  quotidian,  see  p.  167). 
These  various  malarial  fevers  present  certain  clinical  features  in 
common,  which  will  be  stated  here  (see  also  pp.  155  to  157). 
For  further  particulars  regarding  malaria  in  all  its  aspects  the  reader 
is  referred  to  the  book  by  Sir  Ronald  Ross  on  "The  Prevention  of 
Malaria,"  to  the  **  Manual  of  Tropical  Medicine,"  by  Drs.  Castellani 
and  Chalmers,  and  to  the  ** Tropical  Diseases  "  of  Sir  Patrick  Manson. 

Typical  malarial  fevers  consist  of  a  series  of  pyrexial  attacks  which 
recur  at  definite  intervals  of  twenty-four  (quotidian),  forty-eight  or 
seventy-two  hours,  according  to  the  parasite  present  in  the  patient's 
blood.  Each  attack  shows  three  stages,  a  stage  of  rigor,  a  heat  stage 
and  a  stage  of  profuse  perspiration.  Following  on  these  three  stages, 
there  is  an  interval  relatively  or  actually  without  pyrexia.  Then  the 
fever  returns  again.  A  rise  of  temperature,  often  accompanied  by  a 
general  feeling  of  malaise,  may  precede  the  initial  stage  of  rigor.  When 
the  latter  sets  in,  the  patient  feels  intensely  cold,  shivers  violently,  the 
skin  becomes  cold  and  the  features  pinched.     There  may  be  violent 

'  Journ.  Trop.  Med.  and  Hyg.^  xvi,  p.  329. 

^  Annual  Med,  and  Sanii.  Kept.,  Uganda^  for  1913,  p.  80. 


634  THE   ANIMAL   PARASITES   OF   MAN 

vomiting  and  convulsive  attacks  in  young  children.  The  temperature, 
however,  is  really  above  the  normal,  and  continues  to  rise.  After 
about  an  hour,  the  shivering  abates  and  the  heat  stage  succeeds  it. 
The  temperature  rises  rapidly,  even  to  106°  F.  The  patient  becomes 
very  flushed,  the  pulse  is  rapid,  headache  may  be  intense  and  the  skin 
dry  and  burning.  This  stage,  that  causes  acute  distress  to  the  patient, 
may  last  for  one  or  often  three  to  four  hours,  and  then  the  patient 
commences  to  perspire  profusely,  the  clothing  and  bedding  often 
being  saturated  with  sweat.  After  this,  the  fever  rapidly  declines,  and 
when  the  sweating  ceases,  the  patient  may  feel  almost  well  although 
somewhat  languid.  The  sweating  stage  persists  from  two  to  four 
hours,  so  that  the  attack  lasts  as  a  rule  from  six  to  ten  hours.  After 
an  interval  of  one,  two  or  three  days,  a  recurrence  takes  place. 
During  the  early  part  of  the  attack,  especially  at  the  stage  of  rigor, 
there  is  great  splenic  enlargement.  At  first  the  enlargement  disappears 
in  the  interval,  but  in  the  case  of  repeated  attacks  the  spleen  tends  to 
become  permanently  enlarged.  During  malarial  attacks  and  during 
the  intermission  period,  there  is  a  great  increase  in  the  amount  of 
nitrogen  excreted  by  the  kidneys,  while  the  excretion  of  iron  and  bile 
in  the  faeces  is  increased. 

Stitt^  (1914)  points  out  that  it  is  characteristic  of  malignant  tertian 
paroxysms  that  they  set  in  with  chilly  sensations  rather  than  a  frank, 
definite  chill,  and  that  the  fever  is  of  the  remittent  type. 

Plasmodium  malarice  and  P.  %'ivax  rarely  produce  marked  lesions 
in  the  bodies  of  their  hosts,  as  they  sporulate  in  the  circulating  blood 
and  so  do  not  accumulate  in  any  one  organ.  On  the  other  hand, 
Laverania  malarial  (Plasmodium  falciparum)  multiplies  within  the 
internal  organs  of  its  host,  and  consequently  aggregates  or  clusters 
of  the  pai-asites  occur  therein.  The  organ  in  which  most  sporulation 
occurs  suffers  most.  The  liver  is  generally  enlarged,  soft  and  con- 
gested. The  capsule  of  the  spleen  is  tense,  but  the  splenic  consistency 
is  less  than  normal.  The  bone-marrow  is  often  dark  and  congested 
in  the  spongy  bones  and  brownish-red  in  long  bones.  The  blood- 
capillaries  of  the  brain  and  spinal  cord  are  often  filled  or  blocked 
with  sporulating  parasites  and  large  quantities  of  pigment  are  found 
in  these  organs.  Even  if  the  parasites  are  absent,  the  pigment  is 
present  in  the  endothelial  cells.  Pigment  is  found  in  most  organs  of 
the  body. 

Atypical  forms  of  malaria  may  occur  in  v/hich  some  or  all  of 
the  symptoms  are  much  modified.  Irregular  fevers  also  may  be 
produced  by  successive  infections  by  the  same  parasite,  or  by  the 
presence  of  two  different  malarial  parasites. 

'  '*  The  Diagnostics  and  Treatment  of  Tropical  Diseases."     London  :  H.  K.  Lewis. 


SUPPLEMENT  635 

As  regards  the  diagnosis  of  malaria,  according  to  Manson  the 
three  pathognomonic  signs  are — periodicity,  the  effect  of  quinine,  and 
the  presence  of  the  malarial  parasite. 

Trcatineiit. — The  great  specific  for  malaria  is  quinine.  It  attacks 
the  merozoites  or  asexual  generation.  The  drug  can  be  adminis- 
teied  by  the  mouth,  by  the  rectum,  by  intramuscular  injections  or 
by  intravenous  injections,  the  two  latter  methods  being  adopted  in 
serious  infections  or  where  gastric  complications  are  present.  When 
quinine  is  taken  by  the  mouth,  the  more  soluble  acid  salts,  e.^., 
quinine  bihydrochloride  and  bisulphate,  are  better  than  the  sulphate, 
the  form  in  which  quinine  is  usually  sold.  Tablets,  pills  and  cap- 
sules are  convenient  means  of  taking  quinine  but  must  not  be  old 
or  hard,  or  they  may  pass  unchanged  through  the  body.  In  the 
case  of  mild  tertian  or  quartan  malaria,  Castellani  and  Chalmers 
recommend  the  administration  of  a  dose  of  quinine  four  hours 
before  the  sporulation  of  the  parasite  is  due.  Another  modification 
is  to  give  10  gr.  of  quinine  by  the  mouth  in  the  morning  and  a  second 
dose  of  10  gr,  as  above.  In  many  cases  they  give  5  to  10  gr. 
of  the  drug  three  times  a  day.  Administration  of  qumine  per 
reciiim  may  be  useful  but  they  recommend  intramuscular  inocula- 
tion. The  solutions  used  must  be  sterile,  and  the  '^  sterilettes,'' 
small,  hermetically  sealed  vials,  containing  i  grm.  (15  gr.)  or  \  grm. 
(72  &')  o^  quinine  in  solution,  are  recommended.  A  deep  injection 
into  the  deltoid  or  gluteus  muscle  is  usual. 

For  pernicious  infections,  intravenous  inoculation  with  not  less 
than   I  grm.  at  a  time  is  recommended. 

After  the  fever  has  subsided,  the  administration  of  quinine  in 
smaller  doses  must  be  continued  for  some  time,  in  order  to  avoid 
relapses. 

Stitt  (19 14)  writes  that  ''  there  now  seems  to  be  a  tendency  to  use 
the  alkaloid  itself  instead  of  its  salts,  it  having  been  found  that  the 
alkaloid  and  its  very  insoluble  tannate  are  absorbed  from  the  digestive 
tract  equally  as  well  as  the  soluble  salts."  Euquinine  or  ethylcar- 
bonate  of  quinine  contains  81  per  cent,  of  quinine,  but  is  expensive. 

During  malarial  attacks,  constipation  must  not  be  allowed. 
Headache  can  be  relieved  by  cold  applications,  and  perspiration  must 
be  encouraged  in  the  early  stage  by  hot  tea,  warm  lime  drinks,  etc. 
After  bad  attacks,  a  change  to  a  cooler  climate  is  desirable,  but  the 
quinine  treatment  must  not  be  discontinued. 

Preventive  measures  take  two  main  forms,  directed  respectively 
against  the  malarial  parasites  in  man,  and  against  the  mosquitoes 
that  convey  the  parasite  from  man  to  man. 

With    regard    to    man,  houses    should    be  built   away  from   low- 
lying  marshy  ground,  and  kept  free  from  vegetation  such  as  grass  or 
40 


636  THE   ANIMAL   PARASITES   OF   MAN 

brush  which  furnishes  shelter  to  the  mosquitoes.  In  the  tropics,  the 
chief  reservoirs  of  the  malarial  parasites  are  the  native  children,  hence 
European  quarters  should  be  away  from  native  dwellings  as  far  as 
possible.  Mosquito  nets,  having  tw^enty  to  twenty-four  meshes  per 
square  inch,  should  be  used  invariabty,  and  houses  should  be  screened. 
Malaria-conveying  mosquitoes  bite  chiefly  towards  evening.  Quinine 
treatment  for  preventive  purposes  is  important.  A  dose  of  5  gr.  of 
quinine  daily,  with  a  dose  of  10  gr.  on  the  seventh  day  (Castellani),  is 
efiicacious.  Some  workers,  however,  recommend  a  large  dose  (15  gr.) 
on  two  consecutive  days  every  eight  or  ten  days  for  three  months, 
while  others  recommend  10  gr.  twice  a  week.  Celli  administered  3  gr. 
of  quinine  morning  and  evening. 

The  second  line  of  attack  is  directed  against  mosquitoes,  especially 
Anophelines,  on  the  lines  so  well  set  forth  by  Sir  Ronald  Ross.^  The 
accumulation  of  small  quantities  of  water  in  various  vessels,  many  of 
them  unnecessary,  should  be  prevented,  as  Stegomyia  (Culicines)  breed 
in  such  receptacles.  Anophelines  breed  in  small  pools.  All  drinking 
water  and  household  vessels,  water-butts  and  cisterns  must  be 
effectively  screened  with  wire  gauze.  Cesspools,  etc.,  must  also  be 
screened,  and  they,  and  all  collections  of  water,  should  be  oiled  with 
crude  petroleum  sprays  every  week  or  ten  days,  or  fortnight  according 
to  some  workers.  The  petroleum  is  a  good  larvicide  and  suffocates 
the  Anopheline  larvae,  while  its  presence  renders  the  site  obnoxious 
to  the  adult  mosquitoes.  The  amount  of  crude  petroleum  or  kerosene 
will  vary  according  to  the  locality  concerned,  due  regard  being  paid 
to  its  powers  of  spreading  on  the  surface  treated.  Different  authorities 
have  used  different  quantities,  such  as  i  oz.  of  oil  to  i  square  yard 
or  to  15  square  feet.  Others  have  used  i  pint  of  the  petroleum  to 
a  circle  of  20  feet  in  diameter,  while  ^  pint  for  every  100  square  feet 
of  surface  has  also  been  recommended.  The  larvicide  used  so 
successfully  in  Panama  consisted  of  : — 

Average  mixture 
Crude  carbolic  acid  {containing  15  per  cent,  phenol)     ...         300  j;allons 
Caustic  soda         ...  ...  ...  ...  ...  30  lb. 

Resin     ...  ...  .,,  ...  ...  ...         200  ib. 

One  part  of  this  mixture  in  5,000  parts  of  water  containing 
mosquito  larvae  destroys  them  within  five  minutes  ;  i  part  in  8,000 
of  water  kills  larvae  in  thirty  minutes.  Small  fish,  such  as  the 
''millions"  fish,  that  feed  on  the  larvae,  can  be  introduced  into 
collections  of  water  and  are  of  local  service.  Ducks  may  also  act  as 
destroyers  of  larvae.  The  growth  of  water-weeds  and  rank  vegetation, 
tliat  affords  shelter  to  the  larvae,  must  be  prevented  as  far  as  possible. 

Wherever  possible  hollows  should  be  filled  up,  swamps  and  roads 

'  "  The  Preven'irn  of  Malara."     Second  Er  iticn  (191 1).     London  :  John  Murray. 


SUPPLEMENT  637 

should  be  well  drained.  Much  good  has  followed  the  use  of  such 
measures  in  Panama,  Egypt,  British  Guiana  and  other  places.  The 
ideal  conditions  for  malaria  reduction  appear  to  consist  in  a  combina- 
tion of  general  quinine  prophylaxis  with  anti-mosquito  measures. 


YII.— BALANTIDIAN  DYSENTERY. 

This  disease  is  also  known  as  ciliate  or  ciliary  dysentery.  The 
chief  causal  agent  is  Balantidinin  coll.  Others  are  Balantidiuin 
niinutnin,  Nyctof herns  fciba,  etc.  (see  pp.   200-206). 

Balantidiasis  is  insidious  and  is  marked  by  alternate  attacks  of 
diarrhoea  and  constipation  with  vomiting,  while  mucus  is  passed  in 
the  motions,  which  are  foul  smelling.  There  may  be  chronic  ulcera- 
tion of  the  colon.  (Edema  of  the  face  and  limbs  and  ana3mia  may 
occur. 

Treatment  is  at  present  rather  unsatisfactory.  Castellani  and 
Chalmers  state  that  ''the  symptomatic  treatment  for  entamoebic 
dysentery  may  be  tried."  Various  treatments,  more  or  less  empiri- 
cal, by  calomel,  quinine,  carbolic  acid  in  pill  form,  salicylic  acid, 
extract  of  male  fern,  methylene  blue,  iodine  solution,  rice  water  and 
tannin  enemata  are  mentioned  by  Prowazeld  (191 3)  and  by  Seifert. 
E.  L.  Walker^  (1913)  found,  from  experimental  work,  that  organic 
compounds  of  silver,  e.g.,  protargol,  were  most  effective.  Local  treat- 
ment by  large  enemata  of  collargol  or  protargol  seems  to  be  indicated. 
Behrenroth^  (19^3)  successfully  treated  a  Prussian  case  with  thymol, 
given  in  4  grm.  doses  every  two  days,  followed  at  the  end  of  a 
fortnight  by  de-emetinized  ipecacuanha,  given  in  pills  containing 
6  eg.  each,  to  the  number  of  thirty  a  day.  In  about  another  fort- 
night the  symptoms  had  subsided.  The  thymol  checked  the 
diarrhoea,  but  it  was  necessary  to  give  the  de-emetinized  ipecacuanha 
to  kill  off  the  balantidia  still  present,  Phillips  (1915)  also  recom- 
mends thymol.  Ardin-Delteil,  Raynaud,  Coudray  and  Derrieu  (1914) 
found  neither  emetine  hydrochloride  nor  protargol  of  use. 

As  regards  prophylaxis  Walker  states  that  pigs  '*  should  be 
confined  and  not  allowed  to  run  in  yards  and  dwellings."  Behren- 
roth  considers  that  dirty  hands,  for  example,  those  of  farm  workers 
brought  into  contact  with  pigs,  are  probably  the  medium  of  infec- 
tion. The  personal  cleanliness  of  such  persons  is,  then,  of  the 
greatest    importance. 


i 


'  Beihefte  z.  Arch.  f.  Schiffs-  u.  Tropeu-Hyg.^  xvii,  6,  p.  371. 
2  Pkilippiiie  Jl.  Sc,  Sect.  B,  viii,  pp.  1-15,  333-349- 
^   A'ch.  f.    Verdattuu^s  Krankhdten^  xix,  p.  42. 


638  THE   ANIMAL   PARASITES   OF   MAN 


PLATHELMINTHES      (Flat   Worms) 

BY 

J.    W.    W.    STEPHENS,    M.D.,    B.C.,    D.P.H. 

FASCIOLIASIS. 

Fasclola  hepatica. 

The  symptoms  of  disease  evoked  by  Fasclola  hepatica  are  rarely 
observed  in  our  part  of  the  world,  whereas  Kermogant^  states  them, 
to  be  of  frequent  occurrence  in  Tonkin^  ;  the  parasites  are  there 
called  "  Douv-es."  In  our  experience  they  are  only  accidentally 
found  post  inortein  in  a  certain  number  of  cases,  as  no  changes  are 
manifested  during  life  which  would  permit  of  any  conclusion  being 
drawn  as  to  the  presence  of  these  parasites.  In  three  cases  (Bierner,^ 
Bostroem^  and  Sagarra^)  icterus  was  present ;  in  a  fourth  case,  recorded 
by  Duffek/  the  parasites  had  led  to  a  severe  and  acute  distomiasis  of 
the  liver,  combined  with  chronic  purulent  and  ulcerative  cholecystitis^ 
with  purulent  cholangitis  and  dilation  of  the  bile-ducts  and  numerous 
small  abscesses  of  the  liver.  The  total  number  of  flukes  found  in 
these  cases  amounted  to  about  fifty.  The  parasites  passed  from  the 
duodenum  into  the  bile-ducts,  and  first  obstructed  the  flow  of  bile 
and  then  set  up  icterus,  followed  by  cholecystitis  and  cholangitis. 

As  regards  localization  of  the  liver  fluke  in  the  pharynx,  see  p.  242. 

The  treatment  must  be  directed  to  the  principal  symptoms; 
prophylaxis  is  especially  important  in  districts  where  distomiasis  is 
of  frequent  occurrence.  As  the  embryos  live  in  water,  only  boiled 
or  filtered  water  should  be  drunk.  The  attempts  of  Tappeiner^  to 
discover  an  effective  remedy  against  liver-fluke  disease  (liver  rot),  so 
prevalent  among  sheep,  were  unsuccessful. 

Fasciolopsis  buski. 

This  parasite  lives  in  the  intestine,  not  in  the  liver  of  man  ;  it 
produces  bloody  stools  and  typical  symptoms — high  fever  and  a 
condition  of  apathy  (Odhner).^ 

'  Kermogant,  Soc.  vied,  des  HSp.,  February  7,  1905. 

^  [The   distomiasis  of  Tonkin  is  due  to    Clonorchis   sinensis  find    not    to   /'.  hepatica.  — 
J.  W.  W.  S.] 

3  Bierner,  Schwtiz,  Z.eitschr.  f.  Heilk.^  1863. 
--*  Bostroem,  Deutsch.  Arch.f.  Jdiu.  Med.,  1883. 
^  Sagarra,  quoted  by  Dufiek. 
^  Dutifek,  Wiev,  /Jin.  IVcchenschr.,   1902,  xxx. 
■^  Tappeirer,  JMiinch.  vied.  Wccheinchr.^  1900,  I. 
«  Oeilu  er,  CeuhaliLf.  Bald.,  1502,  xxxi. 


SUPPLEMENT  639 

PARAGONIMIASIS. 

Paragonimus    ringeri. 

The  disease  produced  by  the  lung  fluke  is  specially  endemic  in 
Japan,  also  in  isolated  parts  of  China,  Formosa  and  Korea.  The 
fact  that  the  lung-fluke  disease  is  most  frequently  found  in  mountain- 
ous districts  (Katsurada')  is  worthy  of  special  attention.  The  onset 
of  pulmonary  paragonimiasis  is  generally  insidious  (Looss^)  ;  generally 
the  only  symptom  is  a  slight  cough,  occurring  at  first  at  longer,  and 
later  at  shorter  intervals ;  it  is  accompanied  by  the  expectoration  of 
discoloured  sputum,  frequently  blood-stained.  Though  now  and 
then  severe  haemorrhages  result,  up  to  the  present  no  case  has  been 
established  in  which  they  have  been  the  direct  cause  of  death. 

Examination    of   the    thorax  frequently   fails    to    reveal    anvthinr^ 


This    section,     except    for     minor     corrections,     is 
practically  a    translation    of   the    original. 


To  Binder  :    face  p.  638. 


^yinpioms  in  Tnese  tracts.        ' 

The  most  dangerous  locality  is  in  the  brain.  Otani,''^  Inouye,^ 
Yamagiva,^  and  recently  also  Taniguchi,^  have  found  post  mortem 
the  worms  and  their  ova  in  tumours  of  the  brain,  or^  in  areas  of 
softening  in  cases  of  Jacksonian  epilepsy ;  in  Taniguchi's  case  the 
eggs  were  found  in  masses  in  the  inflammatory  areas  of  softening. 
In  the  nineteen  cases  of  paragonimiasis  of  the  brain  collected  by 
Inouye,  the  following  symptoms  were  observed  :  general  convulsions 
on  eight  occasions,  unilateral  convulsions  on  six  occasions,  convul- 
sions with  paralysis  on  the  same  side  and  hemiplegia,  five  times  each ; 

'   Katsurada,  Ziegler^s  Beitr.  z.  path.  Anat.,  igoo,  xxviii. 

^  Looss,  "  Handb.  d.  Tropenkrankh.,"  von  Mense,  iQc;,  i. 

^  Inouye,  quoted  by  Looss. 

^  Scheube,  "  Die  Krankh.  d.  warm.  Lander,"  1896.     ^  Otani,  quoted  by  Looss. 

^  Inouye,  quoted  by  Looss.  "^  Yamagiva,  quoted  by  Looss 

**  Taniguchi,  Arch.  f.  Psych,  ti.  Nervenkrankh..,  xxxviii. 


638  THE  ANIMAL   PARASITES   OF   MAN 


PLATHELMINTHES      (Fiat   Worms) 

BY 

J.    W.    \V.    STEPHENS,    M.D.,    B.C.,    D.P.H. 

FASCIOLIASIS. 

Fasciola  hepatica. 

The  symptoms  of  disease  evoked  by  Fasciola  Jiepatica  are  rarely 
observed  in  our  part  of  the  world,  whereas  Kermogant^  states  them 
to  be  of  frequent  occurrence  in  Tonkin-  ;  the  parasites  are  there 
called  "Douves."  In  our  experience  they  are  only  accidentally 
r.       .1    L  .^j.  v._.  ;,,  ,,  ^^,-fain  nnmhprof   cascs.  as  uo  chauges  are 


prevalent  among  sheep,  were  unsuccessful. 

Fasciolopsis  buski. 

This  parasite  lives  in  the  intestine,  not  in  the  Hver  of  man  ;  it 
produces  bloody  stools  and  typical  symptoms — high  fever  and  a 
condition  of  apathy  (Odhner).^ 

'  Kermogant,  Soc.  vied,  des  HSp.,  February  7,  1905. 

■2  [The   distomiasis  of  Tonkin  is  due  to    ClonorcJiis   sinensis  nnd    not    to   /".  hepatica.  — 
J.  W.  W.  S.] 

^  Bierner,  Schweiz.  Zeitschr.  f.  Heilk.^  1863. 
'1  Bostroem,  Deutsch.  Arch.  f.  kliu.  Med.,  1883. 
■'•  Sagarra,  quoted  by  Dufilek. 
^  Dutifek,  Wien.  klin,  IVcchenschr.,   1902,  xxx. 
"^  Tappeii  er,  A  flinch,  mcd.  IVcchenschr. ,  1900,  1. 
*  Oiihi  er,  Ceiih  all  I.  f.  Bait.,  1502,  xxxi. 


SUPPLEMENT  639 

PARAGONIMIASIS. 

Paragonimus    ringerl. 

The  disease  produced  by  the  lung  fluke  is  specially  endemic  in 
Japan,  also  in  isolated  parts  of  China,  Formosa  and  Korea.  The 
fact  that  the  lung-fluke  disease  is  most  frequently  found  in  mountain- 
ous districts  (Katsurada^)  is  worthy  of  special  attention.  The  onset 
of  pulmonary  paragonimiasis  is  generally  insidious  (Looss^) ;  generally 
the  only  symptom  is  a  slight  cough,  occurring  at  first  at  longer,  and 
later  at  shorter  intervals  ;  it  is  accompanied  by  the  expectoration  of 
discoloured  sputum,  frequently  blood-stained.  Though  now  and 
then  severe  haemorrhages  result,  up  to  the  present  no  case  has  been 
established  in  which  they  have  been  the  direct  cause  of  death. 

Examination  of  the  thorax  frequently  fails  to  reveal  anything 
abnormal.  Inouye^  states  that  the  most  frequently  observed  changes 
consist  in  retraction  of  the  thorax  and  in  a  contraction  of  its  infra- 
scapular  portion.  Scheube*  repeatedly  observed  that  the  one  side, 
presumably  that  which  harboured  the  worm,  moved  less  freely  than 
the  other.  The  physical  changes  are  not  uniformly  spread  over  the 
whole  lung,  but  are  localized.  The  disease  may  come  to  a  standstill 
for  long  intervals  and  then  set  in  again,  lasting  on  the  whole  from  ten 
to  twenty  years.  In  addition  to  paragonimiasis  of  the  lungs,  cysts  are 
frequently  found  on  the  eyelids,  which  occasionally  extend  deeply  into 
the  orbit  and  hinder  the  movements  of  the  eyes.  Post  mortemj  cysts 
the  size  of  hazel  nuts  containing  one,  two,  or  three  adult  worms 
are  found  in  the  lungs,  and  in  addition,  not  uncommonly  there  exist 
pulmonary  emphysema  and  bronchiectasis.  Besides  being  present  in 
the  lungs  and  in  the  eyelids,  the  parasites  have  also  been  found  in  the 
pleura,  the  liver,  the  intestinal  wall,  the  peritoneum,  the  cervical 
glands,  and  in  the  scrotum,  without  actually  occasioning  any  actual 
symptoms  in  these  tracts. 

The  most  dangerous  locality  is  in  the  brain.  Otani,-'*  Inouye,^ 
Yamagiva,^  and  recently  also  Taniguchi,^  have  found  post  mortem 
the  w^orms  and  their  ova  in  tumours  of  the  brain,  or,  in  areas  of 
softening  in  cases  of  Jacksonian  epilepsy ;  in  Taniguchi's  case  the 
eggs  were  found  in  masses  in  the  inflammatory  areas  of  softening. 
In  the  nineteen  cases  of  paragonimiasis  of  the  brain  collected  by 
Inouye,  the  following  symptoms  were  observed  :  general  convulsions 
on  eight  occasions,  unilateral  convulsions  on  six  occasions,  convul- 
sions with  paralysis  on  the  same  side  and  hemiplegia,  five  times  each ; 

'   Katsurada,  Ziegler's  Beitr.  z.  path.  Anat.,  1900,  xxviii. 

■2  Looss,  "  Handb.  d.  Tropenkrankh.,"  von  Mense,  190S,  i. 

^  Inouye,  quoted  by  Looss. 

"*  Scheube,  "  Die  Krankh.  d.  warm.  Lander,"  1896.      '  Otani,  quoted  by  Looss. 

^  Inouye,  quoted  by  Looss.  ^  Yamagiva,  quoted  by  Looss 

^  Taniguchi,  Arch.  f.  Psych,  u.  Nervenkrmikh.^  xxxviii. 


640  THE    ANIMAL    PARASITES    OF    MAN 

in  Taniguchi's  case,  attacks  of  cortical  epilepsy,  choreiform  twitchings 
in  the  right  extremities,  which  gradually  become  athetotic.  The 
following  were  symptoms  of  rarer  occurrence  :  paresis  of  the  right 
upper  extremity,  vertigo,  dementia,  and  amnesic  aphasia,  disturb- 
ances of  vision.  Paragonimiasis  of  the  brain  appears  to  arise  by 
embolism  from  a  primary  pulmonary  lesion. 

The  diagnosis  depends  upon  the  finding  of  ova  in  the  sputa  ;  if 
together  with  ova  in  the  sputa,  cerebral  disturbances  make  their 
appearance,  in  all  probabihty  the  cause  is  the  presence  of  woruis  or 
ova  in  the  brain. 

The  prognosis  of  pulmonary  paragonimiasis  is  favourable ;  on 
the   other  hand,  that  of  cerebral   paragonimiasis   is  very   doubtful. 

The  treatment  of  the  pulmonary  lesion  consists  only  in  paying 
attention  to  the  general  condition  (good  food,  rest,  cough  remedies), 
as  all  attempts  to  destroy  the  w^orms  in  the  lungs  by  means  of  vermi- 
cidal  drugs  administered  internally  or  by  way  of  inhalation  have  so 
far  been  without  result.  The  treatment  of  the  cerebral  lesion  is 
entirely  hopeless.  Trephining  has  been  proposed  for  cases  the  con- 
dition of  which  is  more  favourable,  but  it  has  not  reached  the  stage 
of  performance. 

Prophylaxis  consists  in  general  management :  cleansing  and  if 
need  be  boiling  of  everything  that  is  eaten  or  drunk. 

Clonorchls  sinensis. 

According  to  our  present  knowledge  Cloiiorcliis  sinensis  is  only 
found  in  China  and  Japan  ;  even  the  post-mortem  case  reported  by 
Laspeyres^  was  that  of  an  Asiatic  sailor  who  was  admitted  into  the 
General  Hospital  St.  George,  Hamburg,  in  a  moribund  condition 
with  the  clinical  diagnosis  of  beri-beri.  The  bile-ducts  are  the  usual 
site  of  the  parasite,  though  Katsurada-  has  found  them  also  in  the 
pancreatic  ducts.  In  addition,  it  is  found  not  uncommonly  in  the 
upper  portion  of  the  small  intestine,  especially  in  the  duodenum, 
also,  though  decidedly  rarely,  in  the  stomach.  As  these  sites,  however, 
do  not  afford  the  conditions  necessary  to  life,  they  are  only  found 
here  on  their  way  out  of  the  body  of  the  host. 

The  initial  stage  of  infection  with  this  fluke  generally  runs  a 
symptomless  course  ;  in  proportion  as  the  worms  multiply  the 
following  symptoms  are  manifested  :  First  there  is  a  morbid  sense  of 
hunger  and  irregularity  in  defaecation  ;  at  the  same  time  the  patient 
experiences  a  feeling  of  pressure  and  pain  in  the  epigastrium  and  right 
hypochondrium,  or  just  a  dull  pain.  Pressure  increases  the  pain 
considerably.      The    liver    appears    to    be    enlarged,    sometimes  the 


^  Laspeyres,  "  Dissert.  Kiel,"  1904. 

'^  Katsurada,  Ziegler's  Beitr.  z.  path.  Anat.^  1900,  xxviii. 


SUPPLEMENT  64 1 

enlargement  is  specially  perceptible  over  the  left  lobe  of  the  liver. 
The  patients  maintain  a  proportionately  good  general  state  of  liealth  in 
this  state  for  a  long  time  and  may  hope  to  recover.  In  severe  cases 
there  occurs  copious  and  generally  bloody  diarrhoea,  also  icterus. 
The  next  stages  are  anaemia,  emaciation,  epistaxis,  ascites,  enlarged 
spleen,  and  cachexia,  to  which  the  patient  finally  succumbs.  In 
general  the  course  of  the  disease  is  very  chronic  and  irregular;  m 
winter  and  spring  there  is  generally  improvement,  in  the  summer  and 
autumn  the  patient  gets  worse.  At  post-niorteni  the  bile-ducts  are 
enlarged  and  thickened,  there  is  interstitial  hepatitis  with  enlargement 
of  the  liver,  but  not  to  such  an  extent  as  in  hypertrophic  cirrhosis. 
After  the  initial  enlargement  contraction  of  the  liver  sets  in,  the 
peritoneal  coat  and  capsule  proper  of  the  liver  become  more  or  less 
thickened  in  places.  In  the  pancreas  also  dilatation  and  thickening 
of  the  ducts  occur,  as  well  as  interstitial  inflammatory  processes. 
Obstructions  in  the  portal  circulation  may  lead  to  catarrhal  changes 
in  the  stomach. 

The  diagnosis  is  based  on  the  demonstration  of  ova  in  the  faeces. 

As  a  radical  treatment  is  still  unknown,  consequently  it  can  only  be 
purely  symptomatic.  Prophylaxis  consists  in  the  prohibition  of 
drinking  unboiled  water  or  eating  uncooked  molluscs,  fish,  etc.,  of 
canal  water.  Leaving  the  epidemic  region  may  bring  about  gradual 
recovery. 

BILHARZIASIS. 

Schistosoma  haematobium. 

The  symptoms  of  bilharziasis  are  manifested  chiefly  in  the  urinary 
apparatus,  and  above  all  as  haematuria,  at  the  outset  without  any 
special  troubles.  Later,  however,  it  is  accompanied  by  subjective 
symptoms  in  the  shape  of  feelings  of  pain,  and  of  vague  pains  in  the 
perinaeum  and  lumbar  region,  and  of  burning  in  the  urethra  during 
the  passing  of  urine.  All  the  symptoms  are  usually  aggravated  after 
excesses  in  eating  and  drinking,  and  after  considerable  bodily  exertion. 
Another  condition  found,  but  not  often  mentioned,  is  lipuria  (Stock^)  ; 
the  highest  amount  has  been  2  per  cent,  fat  in  the  urine.  Stock  found 
6  to  20  per  cent,  of  eosinophile  cells  in  ten  cases  examined  by  him. 
They  appear  to  be  increased,  especially  in  the  early  cases ;  Kautsky^ 
also  called  attention  to  the  excessive  degree  of  eosinophilia,  whilst 
Goebel'^  expresses  the  opinion  that  a  specific  toxic  action  on  the 
organism  generally  is  not  developed  in  bilharziasis.  Kautsky*  assumes 
a  toxic  anaemia  as  in  the  case  of  ancylostomiasis.  English  authors 
also  have  called  attention  to  the  eosinophilia  and  to  a  considerable 


'  Stock,  Lancet,  September  29,  igo6. 

'^  Kautsky,   Wien.  klin.  Rundschau^  iQOjj  xxxvi. 

^  Goebel,  Arch.  f.  Schiffs-  u.  Tropen-Hyg.^  I903>  vii. 

^  Kautsky,  Wien.  klin.  Rundschau,  1903,  xxxv 


642  THIi   ANIMAL   PARASITES   OF   MAN 

amount  of  leucocytosis  (Balfour/  Douglas  and  Hardy-).  The 
severe  fonus  occur  almost  exclusively  in  men  ;  symptoms  of  catarrh 
of  the  bladder  make  their  appearance,  vesical  calculi  are  frequently 
found,  whilst  the  formation  of  stone  in  the  kidneys  and  ureters  is 
rare.  Urethral  fistula  occurs  in  bilharziasis,  often  without  stricture, 
and  if  granulations  occur  the  fistula  is  distal  to  them.  GoebeP 
regards  the  bilharzia  fistula  as  a  chronic  burrow^ing  of  pus,  caused  by 
the  irritation  set  up  by  the  ova  as  foreign  bodies  and  consecutive 
restricted  suppuration  ;  and  secondly  as  due  to  the  passage  of  urine 
through  the  defect  in  the  epithelium  or  the  wall  of  the  urethra.  The 
fistulae,  which  are  generally  situated  at  the  neck  of  the  bladder  and  al 
the  membranous  portion,  are  very  tortuous  and  frequently  very 
numerous;  they  often  lie  embedded  in  well-marked  tumours — in  fact, 
in  granulation  tumours  with  marked  inclination  to  excessive  formation 
of  cicatricial  tissue.  The  opening  generally  is  in  the  perineal  and 
scrotal  regions.  In  the  case  of  a  patient,  aged  21,  from  the  Transvaal, 
Kutner*  found  by  cystoscopic  examinations  the  whole  summit  and 
walls  of  the  bladder  covered  with  l^uge  and  small  tumours.  In 
addition  lo  smooth  glistening  tumours,  others  were  more  or  less  dis- 
integrated, and  scattered  large  and  small  cauliflower-like  growths 
occurred.  Like  malignant  growths,  the  tumours  were  inclined  to 
break  down,  the  process  extending  from  within  outwards  towards  the 
surface.  Whether  the  hydrocele  so  frequent  in  Egypt  has  any 
connection  with  bilharzia  is  not  known.  A  frequent  sequela  of 
bilharziasis  is  complete  sexual  impotence   (Petrie^). 

Bilharziasis  of  the  rectum  is  manifested  by  symptoms  of 
dysentery ;  the  repeated  violent  attempts  at  defaecation  lead  in  time  to 
prolapse  of  the  rectum,  which  sooner  or  later  induces  septic  infection 
and  so  death.  In  the  mucosa  of  the  rectum,  polypoid  growths  similar 
to  those  in  the  bladder  are  met  with,  due  to  the  ova  of  the  parasites 
in  the  mucosa  and  submucosa.  In  the  case  of  a  man,  aged  36,  who 
had  lived  for  a  long  time  in  South  Africa,  Burfield*^  found  in  the 
excised  vermiform  appendix  ova  of  Scliistosonia  luvinatohimn ;  he 
assumed  this  to  be  a  gradual  secondary  infection  of  the  appendix, 
whilst  Kelly''  mentions  a  case  of  primary  bilharziasis  of  the  appendix; 
the  eggs  lay  in  the  submucosa  directly  above  the  muscularis. 
Tumours  containing  numerous  ova  are  frequently  found  in  the  region 
of  the  genitalia,  thighs  and  scrotum.  In  one  case  Symmers^  found 
numerous  male  schistosomes  in  the    portal   blood  and  a   copulating 

*  Balfour,  Lancet,  December,  1903.  2  Douglas  and  Hardy,  ibid.,  October,  1903. 
^  Goebel,  CentralbL  f.  d.  Kiankh.  d.  Haim  ii.  Sexualorgane,  xvii. 

*  Kutner,  ibid.,  xvi. 

'"  Petrie,  Brit.  Med.  Joum.,  July,  1903.  ^  Burfield,  Lancet,  February  10,  1906. 

^  Kelly,  quoted  by  Burfield.  ^  Symmers,  Lancet,  January  7,  1905. 


SUPPLEMENT  643 

pair  in  the  left  lung.  Though  schistosome  eggs  have  been  found  by 
some  observers  in  the  lung  tissue,  this  is  nevertheless  the  first  case  in 
which  living  parasites  have  been  found  in  the  lesser  circulation. 
Perliaps  they  got  there  by  way  of  the  external  iliac  vein  from  the 
veins  of  the  bladder  and  rectum. 

In  the  female  sex  bilharziasis  is  incomparably  rarer  than  in  the 
male  and  is  generally  limited  to  haematuria.  Biliiarziasis  of  the 
vagina,  which  takes  the  form  of  an  acute  vaginitis,  is  frequent 
according  to  Milton.^  Horwood^  found  in  one  case  a  polypoid 
tumour  of  the  cervix  uteri,  and  in  the  connective  tissue  of  the  tumour 
Schistosoma  ova,  both  in  masses  and  singly,  It  could  not  be 
established  whether  the  ova  reached  the  vagina  and  thence  the  cervix 
directly,  or  through  the  urine  from  the  bladder. 

Tlie  course  of  the  disease  is  chronic,  and  in  slight  cases,  provided 
fresh  infections  do  not  occur,  is  not  unfavourable;  in  severe  cases 
the  cachexia  caused  by  loss  of  blood,  or  intercurrent  diseases  to 
which  the  patients  easily  succumb — e.g.,  pyelitis,  pyelonephritis, 
pyaemia,  or  ur^emia — lead  to  a  fatal  issue. 

In  regions  in  which  Schistosoma  luvmatohiiun  is  endemic,  or  in 
patients  from  such  regions,  the  diagnosis  is  easy  by  microscopically 
linding  the  eggs  in  the  urine. 

As  regards  the  treatment  of  the  affection  this  much  must  be  said, 
that  so  far  there  is  in  existence  no  certain  remedy.  In  countries 
where  bilharziasis  is  endemic  copaiva  balsam  is  considered  a  specific. 
Kutner  {loc.  cit.),  however,  in  the  case  of  his  patient  who  for  a  long 
time  had  taken  no  inconsiderable  amounts  of  copaiva,  had  no  success 
wortli  speaking  of  to  record.  Urotropin  (three  times  daily,  i  grm.) 
has  similarly  failed,  salol  (075  grm.  several  times  daily)  perhaps  affords 
relief  in  affection  of  the  bladder  (Milton).  Methylene  blue,  oil  of 
turpentine  with  extract  of  male  fern  (Brock^),  or  the  latter  alone  and 
santonin  given  in  small  doses  for  a  week  at  a  time,  in  the  morning,  are 
said  by  Petrie''  to  be  of  value.  Sandwith^  and  Harley^  were  not 
very  successful.  By  way  of  experiment  Kutner  for  some  time  used 
collargol  per  rectum,  proceeding  on  the  assumption  that  this  prepara- 
tion, which  has  proved  of  such  remarkable  service  in  bacterial  infec- 
tion, would  perhaps  render  a  continuance  of  life  difficult  for  the 
bilharzia  worms.  But  this  hope  proved  illusory.  In  order  so  far 
as  possible  to  limit  the  loss  of  blood,  Kutner  regularly  employed 
stypticin  for  long  periods  (three  times  daily,  two  tabloids  of  o'oi  grm.) 


Milton,  quoted  by  Loo?s,   "  Handb,  d.  Tropenkrankh.,"  v.  Mense,  1905,  i,  p.  95. 

2  Horwood,  Brit.  Med.  Journ.,  March  10,  1906. 

3  BxocV,  Jotmi.  of  Path,  and  Bad.,  1893.  ^   Petrie,  loc.  cit. 

^  Sandwiih,  Annal.  of  Surgery,  1904,  xxxix.  "  Harley,  Lancet,  1870. 


644  THE   ANIMAL   PARASITES   OF   MAN 

with  undoubted  success,  in  so  far  that  the  haemorrhages  became 
considerably  less  in  amount.  As  two  patients  in  the  course  of  enteric 
fever  lost  their  haematuria,  Stock  accordingly  recommends  subcu- 
taneous injections  of  Wright's  typhoid  vaccine.  In  the  early  stages 
of  the  rectal  lesion  suppositories  of  iodoform,  ichthyol,  or  narcotics 
might  possibly  be  of  use.  In  the  case  of  urethral  hstulae,  division, 
excision  and  scraping  out  of  the  granulation  tissue  are  recommended  ; 
in  cystitis  with  formation  of  tumours  high  resection  with  curetting 
of  the  tumours  or  their  destruction  with  the  cautery  ;  in  the  case  of 
vesical  calculi,  high  resection,  curetting  the  bladder,  and  then 
drainage.  Tumours  of  the  rectum  must  also  be  removed  by 
operation. 

Prophylaxis  is  important  ;  it  should  be  extended  to  all  modes  of 
using  water,  only  filtered  water  being  drunk,  and  only  boiled  w^ater 
being  used  for  washing.  This  advice  should  be  given  to  tourists  who 
travel  through  the  infected  districts,  and  is  also  recommended  to 
soldiers  and  officials  who  are  despatched  to  the  Colonies.  The  favour- 
able influence  of  change  of  climate  can  only  show  itself  where  fresh 
infections  are  avoided. 

CESTODES. 

GENERAL. 

It  seems  advisable  to  preface  the  section  on  the  Cestodes  with 
some  general  observations  on  the  symptoms  of  disease  provoked  by 
tapeworms,  especially  so  far  as  they  relate  to  the  question  of  toxic 
effects,  and  to  include  the  Nematodes  in  this  discussion.  After  this 
will  follow  a  brief  exposition  of  the  most  important  intestinal  lesions 
causally  connected  with  intestinal  parasites. 

It  is  known  to  every  experienced  practitioner  that  the  different 
intestinal  parasites  can  give  rise  to  a  series  of  nervous  symptoms, 
slight  or  severe,  and  produce,  above  all,  blood  changes — anaemia  of 
the  most  varied  nature,  to  the  extent  of  severe  progressive  anaemia. 
These  symptoms  are  regarded  by  many  authors  as  reflex,  or,  as  in 
the  case  of  ancylostomiasis,  the  main  feature  from  the  loss  of  blood 
caused  by  the  habit  of  life  of  the  intestinal  parasites.  More  fre- 
quently, however,  they  are  regarded  as  toxic  conditions  produced  by 
the  parasites.  In  view  of  this  divergence  of  opinion  there  appears 
to  be  some  advantage  in  defining  clearly  the  present  position  as  to 
the  toxic  action  of  parasites.  Most  interesting  in  this  respect  are 
D ibothriocephalns  latiis  and  Ancylostoiim  diiodenale. 

We  are  indebted  to  the  clinic  at  Helsingfors  for  our  most  detailed 
knowledge   of   bothriocephalus  anaemia.      Reyher^   was  the   first   to 


Reyher,  Deutsch.  Arch.  f.  klin.  Med.,  1886,  xxxix. 


SUPPLEMENT  645 

demonstrate  that  this  parasite  under  certain  circiunstances  can  pro- 
duce a  severe,  progressive  and  sometimes  fatal  anaemia,  which  can  be 
cured,  generally  in  a  surprisingly  short  time,  by  expulsion  of  the 
worm.  Among  the  various  hypotheses  which  have  been  advanced 
as  to  the  mode  of  origin  of  bothriocephalus  anaemia,  the  greatest 
importance  has  been  attached  to  the  assumption  already  mentioned 
by  Reyher,  but  definitely  expressed  by  von  Shapiro/  to  the  effect  that 
Bothriocephalus  latns  produces  a  poison  which  is  absorbed  by  the 
intestine  and  exercises  a  deleterious  influence  on  the  composition  of 
the  blood,  especially  on  the  erythrocytes,  perhaps  also  on  the  blood- 
forming  organs.  This  assumption  is  supported  by  no  slight  number 
of  clinical  and  experimental  investigations.  Podwissotsky^  observed 
severe  blood  changes  in  a  child,  aged  4^,  affected  with  B.  latus.  In 
the  case  reported  by  Pariser^  the  severe  anaemia  in  a  girl  dis- 
appeared fairly  soon  after  expulsion  of  the  worm.  In  that  reported 
by  Schaumann'*  high  fever  accompanied  the  bothriocephalus  anaemia  ; 
he  also  proved  the  haemolytic  properties  of  the  broad  tapeworm. 
The  case  reported  by  F.  Miillef^was  one  of  sev^ere  anaemia.  Also,  in 
the  first  of  the  cases  described  by  Kurimoto^  of  DiplogONOporiis 
grandis  there  were  present  the  same  symptoms  of  anaemia  as  in  the 
case  of  B.  latus.  Meyer'^  observed  severe  anaemia  in  two  youths 
caused  by  5.  latus.  Rosenquist^  has  discussed  the  proteid  metabolism 
in  anaemia.  The  presence  of  B.  latus  produces  in  the  majority  of 
cases  an  increased  proteid  consumption,  to  which  the  blood  change 
generally  corresponds — toxic  anaemia  ;  in  a  further  communication 
he  reports  on  twenty  cases  of  bothriocephalus  anaemia,  nineteen  of 
which  were  cured  by  expulsion  of  the  worms,  while  one  case  proved 
fatal,  and  he  again  emphasizes  the  toxic  properties  of  the  intestinal 
parasites.  In  the  case  reported  by  Bendix,^  that  of  a  girl,  aged  4^^, 
the  anaemia  was  moderate,  whilst  in  the  case  of  Zinn^^  (a  woman, 
aged  30)  the  anaemia  was  so  excessive  that  the  patient  succumbed 
five  days  after  expulsion  of  six  bothriocephalus  heads.  Isaac  and 
van  den  Velden^^  have  established  that  in  the  serum  of  patients  who 
suffer  from  anaemia  due  to  B.  latus,  parasitic  products  are  dissolved,. 


'  von  Shapiro,  Zeilschr.f.  klin.  Med.,  1888. 
^  Podwissotsky, /a/^r(5.  y.  Kinder krankh..,  1S89. 
'^  Pariser,  Deulsch.  vied.   IVochenschr.,  1892. 

^  Schaumann,  Berli.n,  1894,  and  Deiitsch.  med.    PVochenschr.,  1898. 
''  Miiller,  Chariie- Annul.,  xiv. 

6  Kurimoto,  Zeitschr.  f.  klin.  Med.,  xl,  and  Kongr.  f.  inn.  Med.,  Karlsbad,  1899. 
'^  Meyer,  Motint  Sinai  Hosp.  Reports,  1903  and  1904,  iv. 

^  Rosenquist,  Vereinf.  innere  Med.  in  Berlin,  May 6,  1901 ;  and  Zeitschr.  f.  klin.  Med.  xlix. 
^  Bendix,  Deutsch.  Aerzte  Zeitg.,  1904,  i. 
'°  Zinn,  Deutsch.  med.  Wochenschr.,  1903. 
"  Isaac  and  van  den  Velden,  Deutsch.  med.   Wochenschr.,  190  ,  xxvii. 


646  THE   ANIMAL   PARASITES   OF   MAN 

as  shown  by  a  distinct  precipitin  reaction.  Galli-Valerio^  considers 
it  likely  that  toxic  substances  are  secreted  by  the  living  helminthes 
which  produce  a  lowering  or  raising  of  the  body  temperature,  nervous 
disturbances  and  haemolysis.  Tallqvist^  succeeded  in  extracting 
from  B.  latus  a  lipoid-like  body  which  had  a  strong  haemolytic  action. 
The  experimental  anaemia  thereby  produced  differed  in  no  respect 
from  the  severe  chronic  bothriocephalus  anaemia  of  man.  The 
question  as  to  under  what  special  conditions  severe,  and  sometimes 
fatal  bothriocephalus  anaemia  is  developed  is  answ^ered  by  Leichten- 
stern^  and  by  Lenhartz,*  by  the  assumption  that  among  the  Bothrio- 
cephali  some  are  toxic,  that  is,  manufacture  a  poison  which,  when 
absorbed  by  the  host,  produces  a  severe  anaemia. 

Certain  factors  lead  him  to  conclude  that  an  accumulation  of 
poison,  dependent  on  time  and  place,  occurs  in  the  Bolhriocephali. 

In  the  case  of  ancylostome  anaemia,  experience  so  far,  according 
to  Leichtenstern,^'  by  no  means  supports  the  hypothesis  of  a  difference 
in  virulence  of  the  worms  according  to  time  and  locality,  ancylostome 
anaemia  being  rather,  so  far  as  is  known  at  present,  in  all  races  of 
man,  everywhere  and  at  all  times,  simply  and  solely  dependent  on 
the  number  of  ancylostomes,  the  duration  of  the  disease  and — 
within  certain  narrow  limits — on  the  individual  capability  of  resisting 
the  loss  of  blood  and  the  toxic  effect  of  the  parasites.  As  is  shown 
by  a  short  historical  resume  of  the  toxic  action  that  has  to  be  con- 
sidered in  ancylostome  anaemia,  we  must  admit  that  doubtless  here, 
as  in  the  case  of  bothriocephalus  anaemia,  the  toxins  secreted  by  the 
parasites  exercise  a  haemolytic  action,  even  while  admitting  Leichten- 
stern's  contention  that  the  significance  of  the  loss  of.  blood  due  to 
ancylostomes  must  not  be  underrated.  The  toxic  hypothesis  acquired 
a  definite  standing  through  a  series  of  experiments  of  Lussana^  on 
rabbits,  where  he  succeded  in  producing  anaemia  by  injecting  urinary 
extracts  of  ancylostome  patients.  Arslan^  extracted  toxins  from  the 
urine  of  two  ancylostome  patients  and  injected  them  into  rabbits, 
which  thereupon  sickened  and  show^ed  the  same  blood  changes  as 
the  ancylostome  patients.  Retinal  haemorrhages,  so  frequent  in 
ancylostome  anaemia,  which,  according  to  Fischer^  and  Samelsohn,^ 
are  not  due  to  direct  loss  of  blood,  must  also  be  ascribed  to  a  para- 
sitic toxin.     A   further  argument  in  favour  of  the   toxic   hypothesis 

'  Galli-Valerio,  Therap.  Monatsh.,  1905. 

-  Tallqvist,  Zeitschr.  f.  klin.  Med.,  1907,  Ixi. 

^  Leichtenstern,   "  Handb.  d.  Therap.  v.  Pentzoldt.-Stintzlng,"  1898,  2nd  edition,  iv. 

^  Lenhartz,  ibid.,  1903,  3rd  edition,  iv,  p.  607. 

•'  Leichtenstern,  Deutsch.  med.  IVochenschr.,  1899. 

^  Lussana,  Rivista  Clin.  Arch.  ital.  di  din.  Med.,  1890. 

^  Aislan,  Rev.  mens,  des  Mai.  de  VEnfance,  1892. 

*"  Fischer,    Versamml.  d.  ophthal.  GeseUsck.,  1892. 

^  Samelsohn,  ibid. 


SUPPLEMENT  647 

is  furnished  by  the  blood  changes  recorded  by  Zappert,^  Miiller  and 
Rieder,2  Bucklers,^  and  Neusser/  which  must  be  regarded  as  the 
expression  of  toxic  action,  especially  with  reference  to  eosinophilia. 
The  striking  increase  in  proteid  destruction  in  ancylostomiasis 
observed  by  Bohland,^  and  which  ceased  after  the  parasites  had  been 
expelled,  also  gives  additional  support  to  the  assumption  of  toxic 
action.  The  observation  of  Daniels^  also  deserves  consideration  in 
this  connection,  according  to  which  the  presence  of  yellow  pigment 
in  the  liver  and  kidney  cells  is  to  be  attributed  to  blood  destruction 
by  a  verminous  toxin  absorbed  from  the  gut.  Looss^  considers  it 
not  at  all  improbable — in  fact,  almost  certain — that  Ancylostoma,  in 
addition  to  withdrawing  blood,  exert  a  kind  of  toxic  action  on  their 
host. 

Scheube^  attributes  almost  equal  importance  to  the  loss  of  blood, 
the  digestive  disturbances,  and  the  intoxication  induced  by  certain 
metabolic  products  of  the  parasites.  According  to  v.  Jaksch^  ancy- 
lostome  anaemia  is  not-  induced  solely  by  loss  of  blood,  but  by  the 
fact  that  the  parasites  produce  a  ferment  which  has  a  toxic  action 
and  produces  stimulation  in  those  organs  in  which  the  eosinophile 
cells  arise.  The  haemolytic  action  of  ancylostomes  has  frequently 
been  observed  by  Galvagno'^  in  men  employed  in  sulphur  mines. 
According  to  Loeb  and  Smith^^  the  anterior  half  of  the  body  of 
ancylostomes  contains  a  substance  which  probably  causes  anaemia. 
Bauer^^  found  in  the  urine  of  ancylostome  patients  glycuronic  acid, 
which  he  considers  to  be  a  sign  of  metabolic  disturbance  due  to 
parasitic  toxins.  As  has  been  demonstrated  by  Allessandrini,'^  the 
secretio  n  of  glands  in  the  anterior  part  of  the  body  has  a  distinct 
haemolytic  effect  on  the  erythrocytes.  While  the  worm  attaches 
itself  to  the  mucosa  by  means  of  its  teeth,  these  glands  discharge 
their  secretion,  producing  hyperaemia.  The  extravasated  blood  is 
acted  on  by  this  secretion,  so  that  it  can  serve  as  food  for  the 
parasites,  Hynek^^  attributes  eosinophilia  (up  to  20  per  cent.)  to 
a  toxic  action.     Goldmann^^  expresses  a  similar  opinion,  though  he 


'  Zappeit,  IVien.  klin.  IVochenschr.,  1892. 

-  Muller  and  Rieder,  Deiitsch.  Arch.  f.  klin.  Med.,  xcviii. 

3  Bucklers,  Munch,  ined.  Wochenschr.,  1894. 

*  Neusser,  Wien.  klin.   Wochenschr.,  1892. 

^  Bohland,  Milnch.  vied.   Wochenschr.,  1894. 

^  Daniels,  Lancet,  No.  3,725.  '   Looss,  Cenlralvl.f.  Bakt.^  1897. 

"  Scheube,  "  Die  Krankli.  der  warm.  Lander,"  1896. 

"  V.  Jaksch,  Milnch.  med.   Wochenschr.,  1902. 

'0  Galvagno,  Arch,  di  Patol.  e  Clin,  inf.,  1902-1904. 

"   Loeb  and  Smith,  Cenlralbl.  f  Bakt.,  xxxvii. 

'-  Bauer,  Wien.  klin.   Wochenschr.,  1904. 

1^  Allessandrini,  Policlinica,  1904.  ^*  Hynek,  Klin.  Chron.,  1904. 

'^  Goldmann,   Wien.  klin.  Rundschau,  1905. 


648  THE  ANIMAL   PARASITES   OF   MAN 

assumes  that  the  anaemia  is  secondary,  as  the  toxin  of  the  cephaHc 
glands,  as  the  parasites  bite,  penetrates  the  mucosa  and  thence  into 
the  blood,  where  it  dissolves  the  red  blood  corpuscles.  Homani' 
discusses  the  agglutinating  haemolytic  action  of  the  serum  of  ancy- 
lostome  patients.  Whether  Ancylostoma  produce  toxins  and  what 
is  their  nature,  or  whether  the  loss  of  blood  causes  the  anaemia, 
Liefmann^  was  unable  definitely  to  determine  ;  haemolytic  substances 
do  not  appear  to  take  any  part  in  it. 

Berti^  also  is  inclined  to  attribute  the  anaemia  to  metabolic 
products  of  the  ancylostomes;  he  found,  in  fact,  that  a  serum  obtained 
from  a  sheep  (after  subcutaneous  injections  of  the  culture  fluid  of 
ancylostome  larvae)  was  efficacious  in  the  treatment  of  ancylostome 
anaemia.  Peiper'^  likewise  assumes  that  the  parasite  secretes  a  cell 
toxin.  Lobker^  at  the  present  day  still  maintains  that  the  cause  of 
the  disease  must  be  looked  for  really,  if  not  perhaps  entirely,  in  the 
continued  withdrawal  of  blood  by  the  parasites  ;  the  secretion  of 
toxins  by  ancylostomes  has  not  yet,  in  his  opinion,  been  conclusively 
proved.  Except  in  the  case  of  Botliriocepkalus  laUts,  referred  to 
previously,  toxic  action  appears  to  be  of  quite  subordinate  importance 
for  the  other  Cestodes  occurring  in  man — especially  Tcenia  solium  and 
T.  saginata,  which  are  most  frequently  found  ;  thus  Cao^  flatly  denies 
the  presence  of  toxins  in  the  body  of  Taeniae,  while  others,  such  as 
Messineo  and  Calmida,^  Jammes  and  Mandoul,^  consider  they  are 
justified  from  their  investigations  in  concluding  that  Taeniae  contain 
a  specific  toxin.  Messineo'-'  injected,  with  all  bacteriological  pre- 
cautions, extracts  of  Taenia,  dissolved  in  physiological  salt  solution. 
He  invariably  obtained  severe  motor  disturbances  and  frequently 
death.  The  observation  by  Pereira^^  of  a  case  of  chorea  in  which 
rheumatic  and  cardiac  symptoms  were  absent  and  which  after  expul- 
sion of  a  Taenia  was  quickly  cured,  also  favours  the  view  of  a  toxic 
action.  Barnabo,"  however,  was  unable  to  obtain  a  toxin  from  Taenia 
saginata.  Gagnoni,^^  on  account  of  a  marked  eosinophilia  which, 
after  expulsion  of  a  Tcenia  saginata,  fell  within  fourteen  days  to 
I  per  cent.,  assumes  the  formation  of  a  Taenia  toxin.  Dirksen's^^ 
observation  has  reference  to  a  sailor  affected  with  serious  anaemia, 
who,  after  expulsion   of  twelve  pieces  of  Taenia  soliniii,  was  rapidly 

1  Romani,  Gaz.  d.  Osp.,  1904.  ^  Liefmann,  Zeitschr.f.  Hyg.,  1905,  1. 

^  Berli,  Gaz.  d.  Osp.,  1906.  •*  Peiper,  Deutsch.  med.  lVo:henschr.,  1897. 

^  Lobker  and  liruns,  Arb.  atis  dem  kaiserl.  Keichsgesundheitsamt,  1906,  xxiii. 

"  Cao,  Ri/orjua  Med.,  1 901. 

■^  Messineo  and  Calmida,  Centralbl.  f.  Baki.,  xxx. 

**  Jammes  and  Mandoul,  Acad,  des  Sciences,  1904. 

^  Messineo,  Gioru.  med.  del  regio  cserc,  1505. 

'"  Pereira,  Lancet,  September,  1903.  "  Barnabo,  Speiimenfale,  1906,  v. 

"  Gagnoni,  Pediafric,  1903.  ''  Dirksen,  De^iisch.  med.  IVochenschr.,  1903. 


SUITLEMENT  649 

cured.  A  portion  of  the  worm  was  already  breaking  down,  the 
absorption  introducing  into  the  body  highly  toxic  ha^molytic  pro- 
ducts, to  which  the  anaimia  must  be  ascribed.  How  far  the  serious 
disturbances  of  the  nervous  system,  frequently  to  be  observed  in 
cases  of  Hymenolepis  nana,  are  to  be  considered  as  of  purely  reflex 
nature  or  toxic  must  remain  an  open  question  ;  the  same  applies  to 
Dipylidinni  caninnni,  in  which  case  Brandt^  observed  serious  central 
nervous  symptoms.  Caution  is  necessary  in  judging  as  to  any  con- 
nection between  worm  stimulus  and  nervous  symptoms  in  cases  of 
Ascaris  infection.  Peiper^  is  inclined  to  regard  such  nervous 
symptoms  not  as  reflex,  but  rather  as  due  to  a  toxin  contained  in 
the  helminthes,  or  metabolic  in  origin. 

In  cases  of  pernicious  anaemia  when  the  symptoms  disappear  after 
expulsion  of  Ascaruhv  a  toxic  action  must  be  assumed  (Demme^). 
Additional  clinical  observations  do  not,  indeed,  lead  to  any  definite 
conclusion  as  to  the  question  whether  Ascaridce  produce  a  toxin  which 
is  capable  of  causing  more  or  less  injury  either  to  the  nervous  system 
or  to  the  blood,  yet  it  may  be  worth  while  to  give  a  brief  review 
of  this  question.  In  a  case  of  Kutner's,^  that  of  a  girl,  aged  12, 
there  was  a  haemolysis  which  was  cured  after  expulsion  of  twenty- 
four  Ascaridcv.  Attacks  of  opisthotonos  in  a  girl,  aged  16,  ceased 
after  seventy-eight  Ascaridcv  had  been  expelled  (Lutz'^).  Unusually 
serious  disturbances  were  observed  in  a  man,  aged  26,  who  was  rapidly 
cured  by  Drouillard*^  by  the  removal  of  a  great  number  of  Ascarida^. 
The  observations  on  pseudomeningitis  are  of  especial  interest  ;  they 
are  evidently  toxic  in  origin  as  in  the  case  of  Annaratone,^  of  a  man 
who  was  taken  ill  with  gastro-intestinal  symptoms  and  who  died  with 
meningitic  symptoms.  Post  niortcin  the  brain  was  normal,  but  the 
stomach  contained  a  great  coil  of  Ascaridce.  The  cases  of  Delille,^ 
Meriel,^  Papi^^  (the  occurrence  of  Cheyne-Stokes  respiration  has  been 
ascribed  to  the  action  upon  the  centre  in  the  medulla  oblongata  of 
the  products  of  the  Ascaridcv),  and  Taillens^^  related  to  children  in 
which  the  meningitic  symptoms  (meningismus),  partly  serious,  dis- 
appeared with  the  removal  of  the  Ascarida\  Mareo^^  designates  this 
disease  helminthiasis  meningitiformis,  which  exhibits  all  the  symptoms 

*  Brandt,  quoted  by  Pollak  in  Cent^albLf.  Bakt.,  1889,  v. 

^  Pciper,  vide  Seift-rt,  "  Lehrb.  d.  Kinderkrankh.,''  1897,  p.  243. 
^  DeiTime,  vide  Seifeit,  ibid. 

*  Kutner,  Be'l.  klin.   Wochenschr.,  1865. 

•5  Lutz,  Centralhl.  f.  Bakt.  ^  Drouillard,  Journ.  de  Med.,  1900.  xi. 

"^  Annnratcne,  Giom.  vied,  del  regio  esetc,  1900. 

*  Dflille,  Journ.  de  Med.,  May  10,  1907. 

"  Meriel,  Annul,  de  Med.  et  Chir.  inf.,  1900. 

'0  Pani,  Gaz.  d.  Osp.,  1901.  ^^  Taillen?,  Arch,  de  tited.  d'Enf.,  1906. 

'^  Mareo,  Allg.  VVien.  med.  Zeitg,,  1902. 


650  THE   ANIMAL   PARASITES   OF    MAN 

of   meningitis,  but   which    is    caused   by   the   metaboUc   products  of 
A  scar i da'. 

Schupfer/  Duprey-  (observations  in  the  West  Indies,  where  such 
symptoms  are  said  to  be  of  very  frequent  occurrence),  Naab^  (the  flow 
of  w^ater  from  the  mouth  at  night  is  mentioned  as  a  remarkable  fact), 
and  Hammiss*  assume  the  action  of  an  Ascaris  toxin  in  the  clinical 
observations  made  by  them,  mostly  children  with  fever  and  intestinal 
symptoms.  Schupfer  assumes  in  such  cases,  as  he  observed  it  once 
in  a  man,  aged  23,  that  the  disease  termed  Lonibricolse  a  forme  iyphoide 
by  Chauffard  was  due  to  B.  coli  of  marked  virulence  due  to  the 
action  of  the  Ascaridcv.  The  Widal  reaction  was  negative.  Koneff^ 
reports  a  case  in  which  acute  attacks  of  cramp,  trismus,  and  rigidity 
of  the  pupil  disappeared  after  expulsion  of  seven  Ascaridce.  Tetanus, 
as  observed  by  Buchholz*^  in  a  girl,  aged  17,  and  rapidly  cured  after 
expulsion  of  sixteen  Ascaridce,  is  manifestly  rare,  since  only  Rose^ 
mentions  this  as  a  cause  in  his  article  on  Tetanus.  Only  a  few 
experimental  data  exist.  Cattaneo^  could  detect  only  a  very  weak 
toxin  in  Ascaris,  while  Messineo,''  by  injecting  into  animals  extracts 
in  physiological  salt  solution,  invariably  succeeded  in  producing  serious 
motor  disturbances  and  frequently  death.  Interesting  also  are  the 
observations  of  Huber,^^  who,  after  working  with  Ascarida%  suffered 
from  itching  of  the  head  and  neck,  blisters,  swelling  of  the  ear,  con- 
junctivitis, ecchymosis  and  troublesome  palpitation  in  the  head.  He 
consequently  assumes  that  Ascaridcv  can  induce  irritation  by  chemical 
(toxic)  means. 

In  the  case  of  Trichocepliahis  dispar  no  more  than  in  the  case  of 
Ascaris  Inmhricoides  can  we  speak  with  certainty  of  a  toxic  effect,  even 
though  a  number  of  observations  are  available  which  might  justify 
such  an  assumption  as  regards  these  intestin?.!  parasites.  Barth^^  found 
the  brain  normal  in  a  man  who  had  d  ed  with  meningitic  symptoms, 
but  the  intestines  were  full  of  Trichocephalus  dispar;  Gibson^^  records 
the  rapid  cure  of  serious  cerebral  symptoms  after  expulsion  of  Tricho- 
cephalus, so  also  Pascal,'^  Burchhardt^^  and  Rippe.^^^  Moosbrugi^er^^ 


Schupfer,  Gaz.  d.  Osp.  1901.  2  Duprey,  Lamet,  1903. 

^  Naab,  Munch.  meL   Wochenschr.,  1902. 

^  Hammiss,   IVien.  med.   Wochenschr.^  1904,  iii. 

^  Koneff,  quoted  by  Liesen,  "  Dissert.  Eoiin,"  190^. 

6  Buchholz,  Norsk.  Mag.  for  Lcige,  1903. 

^  Rose,  Billroth  and  Piiha,  ''  Chirurgie."         "  Caltaneo,  Arch.f.  Kinder heilk.,  xliv. 

"  Messineo,  Giorn.  vied,  del  regio  eserc. ,  1905. 

'"  Huber,  Deutsch.  Arch.f.  klin.  Med.,  1870,  vii. 

"  Earth,  reported  by  Valleix,  Paris,  1845.  '"  Gibson,  Lancet,  1862. 

'^  Pascal,  quoted  by  Kahane,  KorrespondeJizhl.  f.  Schiveizer  Aerz/e,  1907,  viii. 

"  Burchhardt,  Deutsch.  med.  IVochemchr.,  1880. 

'•'  Rippe,  St.  Petersb.  med.    Wochenschr.,  1907,  i. 

"*  Moosbrugger,  Med.  Corresfondeuzbl.  f   Wurttemberg,  1890. 


SUPPLEMENT  651 

was  the  first  to  draw  attention  to  grave  anaemic  conditions  induced  by 
Trichocephalus,  Morsasca^  and  Becker^  to  progressive  grave  anaemia 
(trichocephalus  anaemia  is  accompanied  by  marked  reduction  of  the 
number  of  red  blood  corpuscles,  of  the  specific  gravity  and  of  the 
haemoglobin,  well-marked  morphological  changes  of  the  red  cell, 
micro-,  macro-,  and  poikilocytosis  and  nucleated  red  cells).  Sandler,^ 
in  his  case  of  a  boy,  aged  11,  who  died  of  anaemia,  assumes  a  tricho- 
cephalus toxin  to  be  the  cause  of  the  disease,  and  Kahane  also  reports 
on  anaemic  conditions  induced  by  Trichocephalus.  Girard,'*  in  addition 
to  symptoms  in  the  gastro-intestinal  tract,  calls  attention  to  those 
arising  in  tlie  blood — anaemia  and  its  sequelae — and  also  to  nervous 
symptoms  :  cerebral  phenomena,  headache,  giddiness,  aphonia,  sym- 
ptoms of  meningitis.  In  a  case  of  Schiller's'^  high  fever  was  present, 
which  probably  set  in  when  the  Trichocephali  present  in  the  gut  in 
great  numbers  commenced  their  parasitic  activity.  Hausmann,^  in 
order  to  explain  the  adaptability  of  Trichocephalus,  assumes  that 
according  to  the  locus  niinoris  resistenticc,  at  one  time  the  reflex  at 
another  the  toxic  action  is  effective,  now  on  one  organ,  then  on 
another  ;  anaemia  being  present  in  most  cases,  frequently  general 
and  local  neuroses  and  cerebral  symptoms  of  various  kinds. 

With  regard  to  the  toxic  action  of  Oxyuris  there  is  only  the  single 
record  of  Hartmann,^  who  noticed  the  disappearance  of  epileptic 
fits  and  psychic  disturbances  in  a  gu'l,  aged  13,  after  the  removal  of 
Oxyuris.  Nervous  disturbances  and  blood  changes  can  but  rarely  be 
attributed  to  Strongyloides.  Silvester^  and  Valdes^  report  on  giddi- 
ness, headache  and  anuria  in  cases  observed  by  them  ;  whether  the 
eosinophilia  recorded  by  Biicklers^^  and  Bruns^^  is  due  to  the  toxin 
of  Strongyloides  must  remain  an  open  question. 

Reference  has  already  been  made  to  the  possibility  that  intestinal 
ciliates  {Balantldluin  coll)  can  also  produce  toxins. 

The  contents  of  echinococcus  cysts  appear  to  contain  a  substance 
only  moderately  toxic,  giving  rise  to  urticaria,  in  a  series  of  cases 
where  the  fiuid  has  escaped  into  the  abdominal  cavity  (during 
puncture).  D.  Muller^^  has  collected  nine  such  cases  out  of  the 
literature,  to  which  may  be  added  six  cases  of  Finsen^^  in  which  the 


I 


'  Morsasca,  abstract  in  Centralbl.  f.  innere  Med.,  1897. 

'^  Becker,  Deutsch.  j?ied.  Wochenschr.,  1902. 

^  Sandler,  ibtd.y  1905.  ^  Girard,  Anual.  deTlnst.  Pasteur,  1901. 

^  Schiller,  Beiir.  z.  klin.  Chir.,  1902,  xxxiv. 

^  Hausmann,  St.  Peter sb.  med.  Wochenschr.,  igoo. 

■^  Hartmann,  Naturforschervers.,  Koln,  1889. 

"  Silvester,  quoted  by  Schliiter,  "Dissert.  Kiel,"  1905. 

"  Valdes,  quoted  by  Schluter,  op.  cit.  '"  Bucklers,  Miinch.  med.  Wochenschr.,  1894. 

"  Bruns,  MiUich.  med.  Wochenichr.^  1907. 
'-  Milller,  D.,  "  Dissert.  Wurzburg,"  1885.  "  Finsen,  quoted  by  D.   Muller. 

41 


652  THE   ANIMAL   PARASITES   OF   MAN 

escape  of  fluid  into  the  peritoneal  cavity  led  to  severely  itching 
urticaria,  which  usually  disappeared  again  after  one  or  two  days. 
On  one  occasion,  nideed,  urticaria  occurred  after  rupture  into  the 
pleural  cavity.  In  the  case  recorded  by  Caffarena^  of  echinococcus 
of  the  right  lobe  of  the  liver,  widespread  urticaria  developed  as  the 
result  of  the  exploratory  puncture.  In  the  case  of  an  echinococcus 
of  the  liver  rupturing  into  the  abdominal  cavity  La  Spada^  ascribed 
the  symptoms  leading  to  death  to  toxic  influence  while  the  peritoneal 
symptoms  w^ere  less  marked.  Eosinophilia  in  hydatid  disease  is  slight 
according  to  the  investigations  of  Bindi^  and  Santucci,*  and  is, 
according  to  Welsh  and  Barling,^  no  certain  sign  of  echinococcus  ; 
it  is  independent  of  the  age,  sex  and  temperature  of  the  patient,  but 
upon  rupture  of  the  cyst  eosinophilia  invariably  sets  in. 

The  question  as  to  the  importance  of  helminthes  in  relation  to 
certain  diseases  of  the  gut  requires  special  discussion,  but  it  concerns 
only  Ascaris  liimhricoideSj  Oxyuris  vermicular  is  y  and  Trichocephaliis 
dispar,  and  the  question  of  appendicitis  first  of  all.  The  entrance  of 
intestinal  parasites  into  the  vermiform  appendix  was  already  known 
to  medical  men  in  the  fifties  of  last  century,  as  is  showai  by  the  works 
of  Merling«  (1836),  Zebert^  (1859),  Platonor^  (1853),  and  Schachtinger^ 
(1861).  Most  of  these  authors  have  considered  intestinal  worms, 
together  with  other  foreign  bodies,  to  be  the  cause  of  appendicitis. 
As  regards  the  part  played  by  these  intestinal  parasites  in  the  etiology 
of  appendicitis,  so  much  discussion  has  taken  place  during  the  last 
few  years  that  it  is  worth  while  to  give  a  resume  of  the  later  views  on 
this  question,  even  though  at  the  outset  it  must  be  admitted  that  the 
matter  is  not  cleared  up.  Bergmann^^  records  a  case  in  which  an 
Ascaris  perforated  the  appendix  and  got    into  the  peritoneal  cavity. 

StriimpelP^  reckons  among  the  symptoms  of  Trichocephalus  the 
possibility  of  a  *'  typhlitis."  On  account  of  the  marked  sensitiveness 
of  the  ileo-caecal  region,  Boas^^  mentions  the  possibility  of  confusing 
it  with  appendicitis.  StilP^  regards  Oxyuris  as  a  principal  cause  of 
catarrhal  affections  of  the  appendix.  Arbore-Rally^*  regarded  severe 
symptoms  of  appendicitis  in  a  boy,  aged  10,  as  due  to  Ascarides. 
In  all  cases  of  appendicitis  Metschnikoff^^  requires  a  microscopical 
examination  to  be  made  for  eggs,  and  considers  treatment  for  worms 

'  Caffarena,  Convers.  din.  Genova,  1902.  ^  l^  Spada,  Gaz.  d.  Osp.,  1904. 

'  Bindi,  ibid.y  1907.  ^  Santucci,  **  Clinica  moderna,"  1905. 

^  Welsh  and  Barling,  Scot.  Med.  and  Surg,  yotirn.,  1907. 


»    >    > 
^0  Bergmann,  Prag.  med.  Wochenschr.^  1890. 


"  Striimpell,  '*  Lehrb.  d.  spez.  Path.  u.  Therap.,"  1894. 

'2  Boas,  Deutsch.  vied.  IVochenschr.,  1895.  ''  Still,  Brii.  Med.  Journ.,  1899. 

^'»  Arbore-Rally,  Arch,  de  Med.  des  En/.,  1900.         '^  Metschnikoff,  Bull,  med.,  1901. 


SUPPLEMENT  653 

carried  out  otherwise  as  a  cause  of  the  frequency  of  perityphHtis. 
Matignon^  does  not  agree  with  this  opinion,  as  in  spite  of  the  extra- 
ordinary frequency  of  intestinal  worms  in  China,  he  has  only  seen 
one  case  of  appendicitis  in  four  and  a  half  years,  and  Des  Barres^ 
expresses  himself  in  similar  fashion.  Out  of  twenty-one  cases  of 
appendicitis  Kirmisson^  discovered  the  ova  of  Trichocephalus  eighteen 
imes  and  the  ova  of  Ascarides  in  three  of  these  cases  ;  in  twelve  cases 
of  enteric  fever  the  examination  for  eggs  was  negative  nine  times. 
Moty^  considers  Oxyuris  to  be  the  sole  cause  in  his  three  cases  of 
appendicitis.  Girard^  ascribes  to  Trichocephali  the  role  of  more 
or  less  septic  foreign  bodies  which  may  bring  about  the  entry  of 
intestinal  bacteria  into  the  appendix,  and  Triboulet^  describes  a 
case  of  appendicitis  which  he  considers  was  due  to  Ascaris.  In 
Morkowitin's''  case  numerous  Oxyuris  had  clearly  caused  the 
appendicitis,  von  Genser^  records  the  case  of  a  boy,  aged  5,  who 
was  operated  on  for  appendicitis,  and  who  passed  through  the  opera- 
tion wound  a  living  Ascaris  on  the  eighteenth  day  after  the  operation. 
In  the  first  case  communicated  by  Schiller^  the  disappearance  of  the 
typhlitic  swelHng  after  the  discharge  of  the  Ascarides  pointed  to  the 
etiological  significance  of  the  parasites,  and  the  same  obtained  in  a 
further  case  published  at  an  earlier  date  by  Czerny  and  Heddaus.^*^ 
In  a  case  abstracted  by  Kaposi^^  Trichocephali  appear  to  have  been 
a  contributory  cause  in  the  production  of  the  appendicitis.  In  a 
further  case  reported  by  Schiller,  where  the  appendix  was  removed, 
was  shown  that  Oxyuris  had  given  rise  to  a  pronounced  appen- 
dicular colic.  In  a  girl,  aged  13,  who  died  from  diffuse  peritonitis, 
Schwankhaus^^  found  that  an  Ascaris  had  perforated  the  appendix. 
Ramstedt^^  found  in  an  extirpated  appendix  a  whole  '^tangle"  of 
Oxyuris,  and  believes  in  the  possibility  of  their  having  provoked  the 
inflammation  ;  he  recommends  an  examination  for  entozoa  before 
the  operation,  without,  however,  after  Metschnikoff's  example,  sub- 
stituting worm  treatment  for  the  operation.  Rostowzeff^^  ascribes 
only  a   minimal    direct   etiological    significance  to   intestinal    worms 

'  Matignon  (abstract),  Munch,  rued.  Wochenschr.^  1901. 

■2  Des  Barres,  Gaz.  des  Hop.,  1903.        ^  Kirmisson,  Annul,  de  Med.  el  Chir,  des  Enf.y  1901. 

4  Moty    abstract),  Mx'inch.  med.  Wochenschr.,  1901,  p.  910. 

G  irard,  Amtal.  de  VInst.  Pasteur,  1901. 
6  Triboulet,  .S'iJc-.  med.  des  H6p.  de  Paris,  1901. 
■^  Morkowitin  (abstract),  Centralbl.  f.  d,  Grenzgebiele,  1902. 
"  V.  Genser,  Wien.  med.  Wochenschr. y  1901. 
0  Schiller,  Beitr.  z.  klin.  Chir.,  1902,  xxxiv. 
^°  Czerny  and  Heddaus,  Und.,  xxi. 

"  Kaposi,  ibid.,  xxviii.  '^  Schwankhaus,  Amer.  Pracl.,  1901. 

'3  Ramstedt,  Deulsch.  med.  Wochenschr.,  1902. 
1*  Rostowzeff,  Russ.  vied.  Rundschau,  1903. 


654  THE   ANIMAL   PARASITES   OF   MAN 

.in  the  origin  of  appendicitis  ;  in  163  cases  he  found  worms  in  three 
-instances.  Wirsaladze^  expresses  himself  in  a  similar  fashion. 
Oppe^  observed  Oxyuris  six  times  in  excised  appendices,  and 
emphasizes  the  opmion  that  in  appendicitis  the  question  of  a  worm 
cure  ought  to  be  taken  into  consideration.  Ascaris  and  Oxyuris,  if 
no  contra-indication  exists,  may  be  expelled,  but  in  the  case  of 
Trichocephakis,  which  frequently  defies  all  expulsive  treatment,  no 
attempt  should  be  made,  but  operation  proceeded  to  forthwith. 
In  a  case  briefly  reported  by  Hanau^  Oxyuris  was  undoubtedly  the 
etiological  starting-point;  in  a  case  of  Galli-Vallerio*  Oxyuris  and 
Trichocephakis.  In  the  opinion  of  Ssaweljews^  in  some  cases  of 
appendicitis,  in  addition  to  other  causes,  intestinal  parasites  play  a 
prominent  part.  The  case  recorded  by  Nason^  is  an  interesting  one ; 
m  this  an  Ascaris  in  the  appendix  became  twisted  with  it  round  a  coil 
of  gut,  causing  obstruction.  Spieler'  argues  against  the  under- 
estimation by  many  authors  as  to  the  part  played  by  intestinal  worms 
in  producing  appendicitis,  although  he  also  does  not  regard  them 
as  a  frequent,  to  say  nothing  of  an  exclusive,  cause  of  the  disease. 
In  a  case  recorded  by  Begonin^  fifteen  Oxyuris  w^ere  found  in  the 
excised  appendix  (the  mucosa  showed  some  ulceration),  and  in  another 
recorded  by  Putnam^  twenty  Oxyuris  were  present  in  the  appendix, 
in  which  there  was  no  evidence  of  any  change.  The  standpoint 
Schilling^*^  takes  is  to  the  effect  that  entozoa  irritate  the  mucosa  and 
can  increase  an  already  existing  inflammation,  but  he  considers  it 
very  questionable  whether  they  can  produce  appendicitis.  Blanchard'^ 
assumes  the  possibiHty  of  a  secondary  infection  arising  from  lesions 
of  the  mucosa  produced  by  helminthes  (Ascaris  and  Oxyuris). 
Moore^^  considers  Trichocephakis  the  excitant  of  the  appendicitis 
in  his  case.  In  a  second  case  of  appendicitis  recorded  by  Auley'^ 
operation  became  unnecessary  owing  to  the  passage  of  the  Ascaridcv. 
Page's^*  case  is  an  interesting  one  ;  it  was  that  of  a  man  who  came 
up  for  operation  with  a  diagnosis  of  appendicitis.  On  incising  the 
abdominal  wall  numerous  Ascarides  were  found  at  the  base  of  the 


'  Wirsaladze,  Bcbnlsch.  Gaz.  Botkina^  1902.         ^  Oppe,  Munch,  vied.  Wochenschr.^  1903. 

^  Hanau,  ibid.,  1903. 

^  Galli-Vallerio,  CentrMl.  f.  Bakt.y  1903,  p.  1094. 

'"  Ssawtljews,  Deutsch.  vied.  Zeifg.,  1903. 

^  Nason,  Jotirn.  Amer.  Med.  Assoc.  ^  1904. 

"'  Spieler,  Wien.  klin.  Wochenschr.,  1904. 

^  Begonin,  Journ.  de  Mid.  de  Bordeaux,  July,  1902. 

^  Putnam,  quoted  by  Spieler.  "o  Schilling,  "  Wurzb.  Abhandl.,"  1905.  v. 

"  Blanchard,  Acad,  de  Med.,  July  3,  1904. 

■'2  Moore,  Brit.  Med.  Journ.,  Augu  t  18,  1906. 

'^  Auley,  ibid.,  i(^o6. 

'^  Page,  New  York  Med.  Journ.,  Januaiy  20,  1906. 


SUPPLEMENT  O55 

wound,  lying  in  cavities  ;  even  after  eight  days  Ascarides  escaped 
from  the  wound.  The  author  assumes  there  was  a  perforation  of  the 
gut  wall  ;  it  is  strange  that  the  worms  were  able  to  exist  a  proportion- 
ately long  time  in  the  muscular  tissue.  Schoeppler^  states  that  there 
is  the  danger  of  an  appendicitis  even  after  the  death  of  an  Oxyuris 
that  has  found  its  way  into  the  appendix.  Oui^  met  with  two 
specimens  of  Trichocephalus  which  had  become  embedded  by  their 
thin  ends  deep  in  the  mucosa.  Frangenheim^  is  not  in  a  position  to 
pronounce  any  opinion  as  to  what  part  intestinal  parasites  play  in  the 
etiology  of  appendicitis.  In  a  case  recorded  by  Kahane'^  many 
Trichocephali  were  found  partly  free  in  the  appendix  and  partly 
embedded  in  the  mucosa  ;  microscopically  appendicitis  was  dia- 
gnosed. At  a  laparotomy  for  salpingitis  Heekes^  found  the  appendix 
elongated,  thickened,  and  containing  about  eleven  Oxyuris  without 
the  mucosa  being  in  any  way  changed.  In  one  case  Andrews^  claims 
Ascarides  to  have  been  the  direct  cause  of  the  appendicitis.  The 
literature  dealing  with  this  question,  so  important  in  our  time,  has 
been  collected  almost  without  any  omissions,  but,  unfortunately, 
no  decisive  opinion  as  to  the  significance  of  parasites  in  appendicitis 
can  be  inferred  from  it.  The  vexed  question  whether  intestinal 
parasites,  especially  Ascaris,  are  able  to  penetrate  the  intestinal  wall  is 
just  as  little  finally  decided.  Leuckart,^  Heller,^  Mosler  and  Peiper,^ 
Henoch/^  Davaine,"  Klichenmeister,'"^  and  Bremser^^  are  opposed 
to  the  idea  that  the  healthy  intestinal  w^all  can  be  penetrated  by 
intestinal  worms,  especially  Ascarides,  whilst  a  whole  series  of  other 
authors  are  of  the  opinion  that  even  the  healthy  intestinal  mucosa  can 
be  perforated.  Among  these  is  numbered  Mondiere,^*  who  is  of  the 
opinion  that  Ascaris,  by  violent  pressure  against  the  mucosa,  forces  it 
so  much  apart  that  it  is  enabled  to  escape  through  the  gap  thus 
formed  into  the  peritoneal  cavity;  this  opinion  is  shared  by 
V.  Siebold.^'^     Rokitansky^^  considers  perforation  of  the  gut  by  Ascaris 

^  Schoeppler,  Centralbl.  f.  Bakt.^  1906. 
2  Oui,  Rev.  prat.  d'Obstet.  ei de  Paed.,  1906. 
•^  Frangenheim,  Samml.  klin.  Vortr.^  1906,  No.  424. 
^  Kahane,  Schweiz.  Korrespondenzbl.^  1907,  viii. 
^  Heekes,  Brit.  Med.  Journ.,  March  16,  1907. 

^  Andrews,  ibid.^  1906.  '^  Leuckart,  "  Die  Parasiten  des  Menschen." 

^  Heller,   *•  Handb.  d.  spez.  Path./'  v.  Ziemssen,  vii. 
'•^  Mosler  and  Peiper,   **  Spez.  Palh.  u.  Ther.,"  v.  Nothnagel,  vi. 
'0  Henoch,  "  Vorlesungen  iiber  Kinderkrankheiten." 
"  Davaine,  "  Traite  des  Entozoaires." 

'^  Kiichenmeister  and  Ziirn,  "  Die  Parasiten  des  Menschen." 
'=^  Bremser,  *'Lebende  Warmer  im  lebenden  Menschen." 
'^  yionAihxe,  Schmidt'' s  Jahrb.,  1840. 

'»  V.  Siebold,  *'  Parasiten"  in  Wagner's  "  Handworterbuch,"  1845. 
'«  Rokitansky,   "  Path.  Anat." 


656  THE   ANIMAL   PARASITES   OK   MAN 

as  at  least  a  rare  occurrence.  Gerhardt^  does  not  doubt  that  the 
worms  can  actively  perforate  the  intestine.  Cases  like  those  of 
Abrault,-  Apostolides,^  Marcus''  (recorded  by  Perls  as  a  valid 
example  of  '*  ascaridophagous"  gut  perforation),  Wischnevvsky,^ 
Galvagno/  Salieri'  certainly  show  that  perforation  of  the  healthy 
gut  wall  cannot  be  denied,  but  at  the  same  time  that  this  occurrence, 
compared  with  the  frequency  of  Ascaridce,  should  be  regarded  as 
exceedingly  rare.  It  is  another  matter  as  to  whether  it  is  possible 
for  the  worms  to  penetrate  an  intestinal  wall  already  diseased, 
especially  when  ulcerated ;  a  whole  series  of  observations  are  in 
favour  of  this.  In  Lini's^  case  (fifty-six  Ascarides  escaped  from  the 
umbilicus  of  a  girl,  aged  7),  in  Gralfe's^  (eighty  Ascarides  escaped 
from  an  inguinal  tumour),  in  Nicolino's^^  (perforation  of  the  intestinal 
wall  with  strangulated  hernia),  in  Liesen's"  (a  living  Ascaris  in  the 
peritoneal  cavity  in  a  woman  suffering  from  a  peritoneal  abscess) — 
in  these  it  is  clear  that  disease  processes  in  the  intestine  preceded  the 
exit  of  the  worms.  In  a  case  described  by  Boloff^^  the  Ascarides 
appear  to  have  produced,  by  forming  a  tight  coil,  necrosis  of  the 
gut  with  perforative  peritonitis.  In  a  case  recorded  by  Lutz^'  the 
perforative  peritonitis  w-as  without  doubt  provoked  by  Ascaris,  and 
in  one  by  Schiller^*  the  Ascaris  had  clearly  gained  access  to  the 
peritoneal  cavity  through  a  gunshot  wound  opening.  In  a  case 
observed  by  Rehn^^  the  w^orm  probably  entered  through  a  gangrenous 
portion  of  the  intestine  in  a  hernial  sac.  Broca^^  is  unable  to  deter- 
mine whether  in  his  case  the  intestinal  perforation  was  primary 
(a  worm  escaped  from  the  abdominal  wound  about  two  months  after 
a  laparotomy  for  suppurative  peritonitis).  The  case  reported  by 
Lutz^^  is  of  special  interest :  it  was  that  of  a  young  man  who  had 
shot  himself  in  the  region  of  the  abdomen,  and  w^ho  died  after  fifteen 
days.  At  the  post-mortem  two  Ascarides  were  found  in  the  pulmonary 
artery ;  they  had  probably  escaped  from  the  intestine,  and  had  gained 
access  to  the  inferior  vena  cava.     Froelich^^  assumes  that  in  his  case 

'  Gerhardt,  quoted  by  Liesen,  *'  Dissert.  Bonn." 

2  Abrault,  quoted  by  Seifert,  •'  Lehrb.  d.  Kinderkrankh." 

'  Apostolides,  Lancet,  1898. 

^  Marcus,  quoted  by  Seifert,   •*  Lehrb.  d.  Kinderkrankh." 

•'■'  Wischnewsky,  quoted  by  Seifert,  ibid. 

^  Galvagno,  Arch,  de  Patol.  et  Clin,  inf.,  1902. 

■^  Salieri,  Rif.  vied.,  1902.  ^  Lini,  Schmidt's  Jahrb.,  1838. 

"  Graffe,  F/otokoll  d.  Ges.  f.  Natur  u.  Heilkiinde,  Dresden,  1853. 
'°  Nicolino,  Clin,  mod.,  1902.  "  Liesen,   "Dissert.  Bonn,"  1904. 

'2  Boloff,  quoted  by  Seifert,  "  Lehrb.  d.  Kinderkrankh." 
'8  Lutz,  Centralbl.f.  Bakt. 

'^  Schiller,  Beitr.  z.  klin.  Chir.,  xxxiv,  p.  200.       ''^  Rehn,  5^1? Schiller,  loc.  cit.,  p.  201. 
"^  Broca,  Rev.  mens,  des  Mai.  de  VEnf.,  1904. 
'■^  Lulz,  IVien.  klin.  Wochenschr.,  190$,  xv. 
'^  Froelich,  Rev.  mens,  des  Mai.  de  I'Enf.,  1897. 


SUPPLEMENT  657 

(a  boy,  aged  11)  the  Oxyuris  were  able  to  penetrate  the  whole 
intestinal  wall,  but  Vuillemin^  considers  this  improbable,  and  is 
more  inclined  to  think  that  the  Oxyurides  penetrated  the  rectum  at 
small  ulcerated  points,  and  thus  gained  access  to  the  perirectal 
connective  tissue.  In  females  Oxyuris  not  only  have  the  power 
of  penetrating  far  into  the  sexual  organs  (Marro^),  and  perhaps  causing 
a  parasitic  endometritis  (Simons^),  but  also  clearly  of  gaining  access 
to  the  peritoneal  cavity  by  way  of  the  tubes,  as  is  to  be  assumed  in 
the  case  recorded  by  Kolb^  (that  of  a  woman,  aged  42,  in  whom  post 
inortein  nodules  were  found  over  the  peritoneum  of  Douglas's  pouch, 
in  which  the  pressure  of  encapsuled  Oxyuris  could  be  demonstrated), 
in  that  reported  by  Chiari^  (adult  Oxyuris  in  Douglas's  pouch)  and 
by  Schneider^  (an  Oxyuris  encapsuled  in  the  pelvic  peritoneum). 
Sehrt's^  case  is  worthy  of  attention  ;  in  this  an  abscess  was  found  in 
the  omentum  with  numerous  Ascaris  ova  in  the  pus  and  a  nodular 
lesion  of  the  peritoneum,  with  Ascaris  ova  encapsuled  in  the  nodules. 
Massive  accumulation  of  Ascarides  may  give  rise  to  a  complete 
occlusion  of  the  gut.  Such  an  occurrence  is  not  so  surprising  as 
might  be  thought  when  one  reflects  that  the  number  of  Ascarides  in 
one  individual  may  amount  to  several  hundreds.  For  instance,  one 
boy  evacuated  within  a  single  day  600  Ascarides  (Fauconneau- 
Dufresne^)  and  within  three  years  5,126  worms.  In  the  case 
recorded  by  Tschernomikow^  a  boy,  aged  2^,  evacuated  during  a 
day  208  worms,  partly  through  the  stomach,  partly  through  the 
intestine.  Coil-formation  of  such  masses  of  Ascarides  renders 
possible  not  only  constipation,  but  also  complete  obstruction  with 
symptoms  of  ileus,  as  shown  by  the  five  cases  quoted  by  Mosler 
and  Peiper,^^  as  w^ell  as  from  observations  made  by  Raie,^^  Schulhof,^^ 
Rehberg,^3  Rocheblave,^*  Heller,^^  Leichtenstern,^^  Huber,^^  and 
VVilms.^^  In  two  cases  of  Black^^  and  Parkinson^*^  the  intestinal 
obstruction  was  caused  by  a  coil  of  tapeworms. 

In  the  earlier  history  of  medicine  the  helminthes  played  a  great 
part  as  the  excitants  of  many  intestinal  diseases  and  of  enteric 
as  well.  Even  if  to-day  they  no  longer  be  regarded  as  such,  the 
conception  that    they   represent   the  predisposing  factor  in   typhoid 

^  Vuillemin,  Centralbl.  f.  Bakt.,  1902,  ^  Marro,  Arch,  per  le  Set.  med.,  1901. 

»  Simons,  Centralbl.f.  Gyndk.,  1899.  ^  Kolb,  Ceniralbl.f.  BakL,  1902. 

'"'  Chiari,  Prag.  me  J.  Wochenschr.,  1902.  ^  Schneider,  Centralbl.f.  Bald.,  1904. 

"^  Sehrt,  Beitr.  z.  klin.  Chir.,  li.  ^  Fauconneau-Dufresne,  quoted  by  Seifert. 


9  Tschernomikow,  quoted  by  Seifert. 

"  Raie,  Lancet,  1899. 

'3  Rehberg,  **  Dissert.  Konigsberg,"  1907. 

'^  Heller,  loc.  cit. 

'^  Huber,  quoted  by  Rehberg. 


°  Mosler  and  Peiper,  loc.  cit. 
2  Schulhof,  Milne h.  med.  Wockemchr.,  1903. 
'  Rocheblave,  Gaz.  des  Hdp.,  1898. 
6  Leichtenstern,  "  Ziemssen's  Handb.,"  vii. 
Wilms,  Deutsch.  Zeitschr.  f.  Chir.,  xlvi. 


'9  Black,  Brit.  Med.  Joiirn.,  1872.  ^o  Parkinson,  quoted  by  Rehberg. 


658  THE   ANIMAL   PARASITES   OF   MAN 

infection  through  the  injury  they  inflict  on  the  mucosa  (Guiart/ 
Blancliard,^  Vivaldi  and  Tonello^)  must  not  be  summarily  rejected. 
Vivaldi  and  Tonello  found  helminthes  in  80  per  cent,  of  their 
typhoid  patients,  numbering  among  these  Trichocephahis  dispar, 
Oxyiiris  vennicularls,  Ancylostoma  diiodenale,  and  Ascaridct.  The 
report  of  Leuckart"^  is  here  worth  citing,  to  the  effect  that  Thiebault 
never  failed  to  find  Trichocephalus  in  his  cholera  patients  at  Naples. 
Blanchard^  goes  so  far  as  to  express  the  desire  that  in  every  febrile 
affection  of  the  intestine  an  anthelmintic  treatment  with  thymol 
should  be  undertaken  as  early  as  possible,  even  before  learning  the 
results  of  serum  diagnosis. 

The  lesions  of  the  liver  and  pancreas  due  to  Ascaridcv  are  briefly 
discussed  in  the  chapter  on  Ascariasis  (p.  687). 

A  discussion  of  the  intestinal  helminthes  from  the  clinical  and 
therapeutical  point  of  view  follows  these  general  considerations. 

Dibothriocephalus  latus. 

From  what  is  known  as  to  the  development  of  Dibothriocephalus 
latus,  the  way  by  which  man  is  infected  is  self-evident  :  infection 
can  only  take  place  through  the  ingestion  of  insufficiently  cooked 
fresh-water  fish  (pike,  burbot,  perch,  grayling  and  vendace) ;  what 
degree  of  temperature  is  necessary  to  kill  the  larval  forms  is  still 
unknown.  Dibothriocephalus  latus  lives  in  the  small  intestine  of  man, 
alone  or  in  some  numbers,  frequently  also  together  with  Taenia  solium. 
The  proglottides  are  passed  always  united  in  large  pieces,  the  ova  are 
deposited  through  the  uterine  pore,  while  the  worm  is  still  in  the 
intestine,  so  that  they  are  easily  found  in  the  faeces.  The  proglottides 
are  so  characteristic  that  they  cannot  be. confused  with  those  of  other 
species.  In  reference  to  whether  age  or  sex  is  spared  by  Z).  latus, 
it  is  not  possible  to  make  any  definite  statement,  especially  so  far 
as  the  endemic  area  is  concerned,  whether  a  person  resides  n  it 
continuously  or  visits  it,  so  long  as  his  habit  of  life  is  in  accordance 
with  those  of  the  country.  Bendix^  certainly  emphasizes  the  fact 
that  early  childhood  is  as  a  rule  immune  :  his  case  was  that  of 
a  child,  aged  4^  years. 


*  Guiart,  Compt.  rend.    Soc.  de  BioL,  Paris,  March  i6,  1901. 

2  Blanchard,  A^-ch.  d.  Par.,  1901. 

^  Vivaldi  and  Tonello,  Gaz.  d.  Osp.,  October  29,  1905. 

4  Leuckart,  quoted  by  Kahane.  s  Blanchard,  Acad,  de  Med.,  Ocloler  18,  1904. 

fi  Bendix,   Verein  f.  inncre  Med.,  Berlin,  June  16,  1902, 


SUPPLEMENT  659 

Sparganum   mansonl. 

According  to  our  present  knowledge  (Miyake^)  the  disease  occurs 
almost  exclusively  in  China  and  japan.  On  the  main  island  it  occurs  in 
all  districts,  though  rarely  under  observation.  It  is  especially  frequent 
in  the  neighbourhood  of  Kioto  and  Osaka;  these  places  are  very 
near  together,  and  between  them  there  is  mutually  active  intercourse, 
so  that  taken  together  they  may  be  regarded  as  one  district  infested 
by  this  worm  disease.  As  regards  localization  in  the  body,  there 
appears  to  exist  a  certain  predisposition  for  definite  regions,  for 
instance,  the  eye  and  genito-urinary  tract.  In  some  cases  the  parasite 
manifested  the  peculiarity  of  wandering  about  the  body  and  of 
appearing  at  certain  favourite  points  (muse,  quadriceps  femoris) 
(Hashimoto^).  Most  patients  complain  more  or  less  of  the  onset  of 
attacks  of  pain  and  of  sensitiveness  to  pressure.  In  those  cases  in 
which  the  patients  evacuated  the  worm  during  micturition,  the 
symptoms  were  variable ;  sometimes  there  was  tenesmus  of  the 
bladder,  sometimes  pains  in  the  inguinal  region,  sometimes 
haematuria.  None  of  these  troubles  is  characteristic  of  the  disease, 
and  does  no  more  than  represent  the  symptoms  that  follow  a 
mechanical  irritation  that  any  kind  of  foreign  body  may  produce. 
Besides  the  onset  of  attacks  of  pain,  swelling  of  the  regions  affected, 
if  superficial,  may  often  be  recognized,  when  a  superficial  diffuse 
soft  tumour  can  be  felt  which  often  gives  pseudo-fluctuation.  Some- 
times a  peculiar  crackling  can  be  detected  internally,  as  in  the  making 
of  a  snowball.  During  the  further  course  an  abscess  not  infrequently 
forms  around  the  worm.  When  the  situation  of  the  worm  is  super- 
ficial, *' an  inflammatory  tumour  with  a  tendency  to  migrate"  is 
stated  by  Omi^  to  be  an  important  diagnostic  sign.  That,  however, 
is  not  always  the  case,  as  the  observation  made  by  Inoye"^  shows.  It 
would  be  better  to  add  to  this  sign  the  onset  of  paroxysmal  pain  and 
the  temporary  change  in  volume  of  the  tumour.  When  once  the 
parasite  is  removed,  the  wound  heals  just  as  satisfactorily  as  any 
other  fresh  wounds  made  at  operation. 

Dipylidium  caninum  {Tcenia  ciiaunerina). 

This  species  belongs  to  parasites  of  rare  occurrence.  Up  to  the 
year  1905  Bollinger^  collected  thirty-six  cases  from  the  literature, 
twenty-nine  of    which  were    children  and  seven  adults.     Since  then 

'  Miyake,   Mitleil.  aus  d.  Grenzgebieie,  1904,  xiii. 

-  Hashimoto,  quoted  by  Miyake.  ^  Omi,  Jji-Shinshi^  Tokio,  1898. 

^  Inoye,  ibid.^  1897. 

^  Bollinger,  Deutsch.  Arch.f.  klin.  Med.j  1905,  Ixxxiv. 


66o  THE   ANIMAL   PARASITES   OF   MAN 

some  further  cases  have  come  to  liglit,  so  that  the  number  now 
observed  amounts  to  ninety,  and  among  them  only  eight  adults. 
The  youngest  child  was  6  weeks  old  (KohF),  in  which  the  first  pro- 
glottides were  passed  when  the  child  was  40  days  old.  This  pre- 
ponderating occurrence  in  children  is  clearly  connected  with  the 
close  intercourse  between  children  and  dogs,  and  also  cats.  Bollinger 
believes  that  D.  canimim  in  reality  occurs  more  frequently  in  adults 
than  has  hitherto  been  supposed.  In  addition,  it  must  be  mentioned 
that  this  species  is  quite  unknown  to  many  physicians,  and  is 
occasionally  confused  with  Tcenia  soliutn.  One  notices  almost  daily 
a  large  quantity  of  cucumber-seed-like  bodies,  reddish  or  whitish- 
grey,  about  I  cm.  long  and  2  mm.  broad,  discharged  with  the  stools. 
Lindblad'^  remarks  that  these  bodies  have  lively  movements,  that  they 
perish  rapidly  in  fresh  water,  and  become  white  and  smooth.  These 
Cestodes,  in  isolated  cases,  are  parasitic  in  the  intestine  in  large 
numbers.  Sonnenschein^  expelled  four  fragments  in  the  case  of  a 
boy,  aged  4  months  ;  Asam^  three  fragments  in  the  case  of  a  child,  aged 
19  months  ;  and  Zschokke'^  as  many  as  five  or  six  in  that  of  a  boy, 
aged  4.  They  do  not  always  produce  such  striking  symptoms  as 
occurred  in  Pollak's  case.^  In  other  cases  gastro-intestinal  dis- 
turbances with  or  without  fever  (Kriiger^),  emaciation  (Zschokke), 
or  even  nervous  symptoms  of  central  origin  in  the  form  of  con- 
vulsions (Brandt^)  have  been  observed.  From  the  nature  and  mode 
of  infection  children  must  be  kept  from  close  contact  with  dogs  and 
cats  as  much  as  possible  lo  ensure  prophylaxis.  The  appropriate 
treatment,  as  it  mainly  affects  children,  deserves  special  mention, 
whilst  the  expulsion  of  the  remaining  Cestodes  may  be  described  in 
this  connection.  Among  the  drugs  one  may  mention  flor.  kousso 
I'D  grm.,  pulpa  tamar.  depur.  2  grm.,  syrup  of  sugar  50-0  grm.,  one- 
third  to  be  taken  every  hour  (Lindblad).  Kamala  appears  to  have  no 
effect,  although  Huber^  recommends  it  in  small  doses  according  to 
age  from  0*5  to  30  grm.  He  warns  against  Filix  mas  preparations, 
which  otherwise,  even  in  early  childhood,  vuider  careful  dosage  gives  the 
best  results.  Young  children  are  given  i*o  to  2*0  grm.  extr.  fil.  maris, 
with  mint  syrup  or  raspberry  syrup  30*0  grm.,  in  the  morning  twice  an 
hour  by  the  mouth,  or  i-q  grm.  extr.  fil.  maris  is  mixed  with  syrup 


^  Kohl,  Munch,  med.  Wochenschr.,  1904.  -  Lindblad,  Hygiea,  xlv. 

^  Sonnenschein,  Milnch.  med.  IVochenschr.,  1 903. 

^  Asam,  Afilnch.  med.  Wochensch}-.^  1903. 

^  Zs~hokke,  Cenfralbl.  f.  Bakt.,  1905. 

"  PoUak,  Wien.  klin.  Wochenschr.,  1907. 

■^  Kriiger,  St.  Petersb.  nied.  IVochenschr.,  1887. 

»  Brandt,  Cenfralbl,  f.  BakL,  1889. 

•"  Huber,  suppl.  to  Asam,  Munch,  med.  IVochenschr.^  1903. 


SUPPLEMENT  66l 

of  mint,  and  given  by  means  of  a  stomach  tube  (Rosenberg^).  A  few 
hours  afterwards  a  mild  laxative  may  be  taken — one  to  two  table- 
spoonfuls  of  aqueous  tincture  of  rhubarb  (Asam) — or  an  enema  may 
be  given.  In  a  case  reported  by  Sonnenschein  decoction  of  pome- 
granate root  had  no  effect,  as  it  was  vomited  up. 

Hymenolepis  nana. 

This  species,  very  rare  in  Central  and  Northern  Europe,  inhabits 
the  small  intestine,  especially  of  children  ;  it  burrows  very  deeply  into 
the  mucosa.  Not  uncommonly  several  thousand  have  been  found  in 
one  case  (Nicolo,^  E.  Stoerk  and  HaendeP).  It  is  remarkable  that 
these  Cestodes  have  been  found  so  frequently  post  inoiiem  and  after 
vermifuges  given  for  other  reasons.  Thus  the  clinical  symptoms 
must  often  be  very  indefinite  (Stoerk  and  Haendel),  so  that  one 
may  assume  that  only  a  slight  percentage  of  cases  of  Hynienolepis 
liana  come  under  observation  and  are  published  as  such.  On  the 
other  hand,  it  is  certainly  conceivable  that  with  the  large  number  of 
parasites  that  frequently  occur  in  one  individual  a  whole  series  of 
symptoms,  in  part  quite  severe,  are  capable  of  being  produced. 
These  are  partly  symptoms  of  intestinal  catarrh,  consisting  of 
abdominal  pains,  constipation,  alternating  with  attacks  of  diarrhoea, 
perverse  appetite,'  and  boulimia,  abdominal  pains  of  a  cramp-like 
nature,  followed  by  emaciation,  headache,  sleeplessness,  pallor, 
lassitude,  and  in  part  nervous  symptoms — epileptiform  attacks 
without  loss  of  consciousness,  weakness  of  memory,  melancholia, 
irregular  febrile  attacks  (Lutz^).  Possibly,  too,  Hymenolepis  nana 
infects  the  urinary  organs,  producing  true  chyluria  (Predtetschensky^). 
Stoerk  and  Haendel  are  inclined  to  think  that  this  species,  unlike 
other  Cestod.es  parasitic  in  man  and  domestic  animals,  needs  no 
intermediate  host  for  its  development,  and  that  the  larval  forms 
(cysticercoid)  live  in  the  same  host  as  the  adults.  The  diagnosis 
is  based  on  the  demonstration  of  ova  in  the  stools.  As  far  as 
expulsion  of  this  Cestode  is  concerned,  santonin,  kamala,  kousso 
flowers  and  thymol  appear  to  have  no  effect  of  importance  ;  whilst 
extract  of  male  fern,  recommended  by  Grassi^  as  a  result  of  his 
considerable  and  successful  experience,  has  been  given,  with  the 
result  that  the  worms  really  are  expelled,  and  that  after  the  treatment 
neither  worms  nor  ova  are  any  longer  demonstrable  in  the  stools  of 
patients.      In   his  cases  of   chyluria    Predtetschensky   prescribed    ol. 

1  Rosenberg,  Ges.  f.  innere  Med.,  February  i6,  1904.  ^  Nicolo,  Gaz.  d.  Osp.,  1904. 

^  Stoerk,  E.,  and  Haendel,  Wien.  klin.  IVockenschr.,  1907,  xxix. 
*  Lu'z,  Centralbl.f.  Bakl.,  1894.  ■'  Predtetschensky,  Zeitschr.  f.  klin.  Med.,  xl. 

"  Grassi,  Centralbl.  f.  Bak:.,  1887. 


662  THE   ANIMAL   PARASITES   OF   MAN 

terebinth.  20  drops  three  times  daily  for  a  fortnight,  then  acid,  galhc. 
0*5  grm.  three  times  a  day  for  two  days,  then  i*o  grm.  three  times  a 
day ;  the  urine  became  clear,  but  whether  permanent  cure  resulted 
remained  doubtful. 

Hymcnolcpis  diminiita,  H,  lanceolata,  Davainea  asiaiica,  and  D. 
madagascarensis  possess  no  actual  clinical  interest ;  with  regard  to 
the  latter  it  need  only  be  pointed  out  that  Bordier^  in  studying  a 
case  of  chyluria  found  this  species  in  the  kidneys  of  a  person  in 
Madagascar. 

Taenia  solium. 

Tceiiia  solium  inhabits  the  small  intestine  of  man ;  single 
proglottides  or  whole  worms  may  get  into  the  abdominal  cavity  and 
the  bladder  through  fistulae,  and  penetrating  the  abdominal  wall 
escape  outwards  or  become  discharged  with  the  urine.  Symptoms  of 
intestinal  stenosis  are  certainly  very  rare,  as  in  the  case  recorded  by 
Steinhaus^  of  a  child,  aged  9,  the  stenosis  ceasing  after  the  expulsion 
of  the  segments.  The  usual  position  of  the  worm  in  the  small 
intestine  is  with  the  head  closely  adherent  to  the  mucosa  and  the 
proglottides  lying  along  the  intestine  ;  from  time  to  time  portions  are 
discharged  with  the  faeces  per  rectum.  Its  position  can  also  be 
reversed,  and  the  proglottides  in  the  gut  become  thus  discharged  by 
vomiting. 

The  diagnosis  depends  upon  the  proglottides  being  generally 
discharged  in  pieces  in  the  stools,  or  eventually  an  examination  for 
eggs.  Larval  infection  (Cysticercus  celhdoscv)  occurs  also  in  man 
through  auto-infection  or  through  food. 

Cysticercus  cellidosce  of  the  skin  and  subcutaneous  tissue  occurs 
very  seldom  singly ;  as  a  rule  they  are  found  in  hundreds  and 
thousands  in  the  same  individual.  They  occur  in  different  parts  of 
the  body,  especially  on  the  flexor  surfaces  of  the  extremities  (generally 
symmetrically),  small  globular  swellings,  the  size  of  a  pea  or  a  hazel 
nut,  smooth,  of  a  tough  cartilaginous  consistence,  fairly  movable 
under  the  skin,  in  the  muscles  less  so.  They  never  degenerate  or 
cause  the  surrounding  skin  to  lose  its  colour.  It  is  an  interesting 
fact  that  in  the  case  described  by  Possell^  nodules  on  the  face, 
namely  in  the  neighbourhood  of  the  left  cheek  and  behind  the  left 
ear,  reformed.  The  following  are,  according  to  Posselt,  characteristic 
for  cutaneous  tumours  due  to  cysticerci  :  (i)  the  position  in  the 
subcutaneous  connective  tissue  (and  almost  always  simultaneously 
in   the   muscles)  ;     (2)    the   approximately   equal    size   and   regularly 

/  Bordier,  quoted  by  Predtetschensky,  loc.  cit.,  p.  95. 
^  Steinhaus,  Deutsch.  med.  Wochenschr.^  1903. 
8  Posselt,  Wien.  klin,   IVochenschr.y  1899. 


SUPPLEMENT  663 

rounded  oval  form ;  (3)  the  peculiar  density,  almost  reminding  one 
of  cartilage  in  its  hardness  and  the  sensation  of  tightly  distended 
thick-walled  bladders;  (4)  proportionately  slight  mobility;  (5)  with 
painlessness,  absence  of  any  cutaneous  reaction  (hyperaemia  or 
swelling  of  the  skin  or  pigmentation).  The  very  gradual  appear- 
ance generally  of  the  tumours  supports  the  diagnosis,  and  in 
addition  to  this  evidence  we  may  emphasize  the  preponderating 
liability  of  the  upper  part  of  the  body  to  attack  and  the  symmetrical 
arrangement  of  the  nodules.  Cutaneous  and  muscular  cysticerci 
cause  the  most  varied  symptoms,  sensory  disturbances,  abnormal 
sensations,  depression  and  a  feeling  of  weariness  whenever  the  diseased 
parts  are  moved,  weakness  in  the  lower  extremities,  pains  in  the  course 
of  the  sciatic  nerve,  in  addition  to  those  which  simulate  cramp  in  the 
calves,  numbness  in  the  hands,  pains  upon  their  being  moved.  In 
the  case  of  a  cysticercus  situated  in  the  elbow-joint,  painful  dragging 
sensation  in  the  course  of  the  ulnar  nerve  persisted.  In  other  cases 
the  arm  was  almost  paralysed,  or  it  could  not  be  completely  extended ; 
stiffness  and  bending  of  the  little  finger  were  noticed.  Cysticerci  of 
the  gluteal  muscle  cause  trouble  upon  sitting  and  upon  defaecation. 
Remittent  unilateral  headaches  were  present  in  the  case  of  a  cysti- 
cercus of  the  region  of  the  right  eyebrow  ;  pains  of  a  neuralgic 
character  radiated  from  the  diseased  temporal  region.  The  cysts  may 
be  inflamed  and  may  suppurate ;  this  especially  happens  in  the  case 
of  solitary  cutaneous  and  muscle  cysticerci.  The  best  treatment 
consistS'in  puncture  of  the  cysts  with  a  Pravaz  syrmge  and  subsequent 
injection  of  a  drop  of  i  per  cent,  sublimate  solution.  Tincture  of 
iodine  has  similarly  been  proposed  (Wolff^).  Frangenheim^  re- 
commends early  extirpation  (this,  however,  only  in  the  case  of 
solitary  cysts).  Pelagutti^  believes  that  in  his  case  diminution  in 
the  size  of  the  cysts  was  obtained  by  the  use  of  anthelminthic  remedies 
continued  over  a  long  period  combined  with  potassium  iodide  and 
calcium  salts  (internally).  Cysticercus  is  very  rarely  found  in  the 
tongue  ;  there  the  w^orms  generally  lie  in  front  of  the  sulcus  terminalis, 
corresponding  to  the  middle  of  the  tongue,  according  to  Glas.^  In 
the  case  recorded  by  Gaetano^  (a  boy,  aged  10)  there  was  a  nodule 
on  the  left  side  of  the  tongue  which  grew  very  rapidly  till  it  reached 
the  size  of  a  nut  ;  it  was  embedded  in  the  muscle  and  covered  over 
by  normal  mucosa.  Cysticerci  are  just  as  rare  in  the  pleurae,  in  the 
lungs,  in  the  intestinal  submucosa,  in  the  submucosa  of   the  small 

'  Wolff,   "  Lesser's  Encyclop.  d.  Haut-  u.  Geschlechtskrankh.,"  1900. 

^  Frangenheim,  Volkni.  klin.  Vortr.,  No.  424. 

^  Pelagutti,  Giorn.  ital.  delle  nial.  vener.,  1900. 

^  Glas,  IVien.  kliit.   IVochenschr.y  1905. 

^  Gaetano,  Gioin.  int.  delle  Set.  med,^  1904. 


664  THE   ANIMAL   PARASITES   OF  MAN 

intestine,  in  the  mesenteric  glands,  in  the  liver,  pancreas,  spleen  and 
kidneys,  in  the  mamma,  in  the  heart,  in  the  bones  and  in  the  great 
vessels  (Huber^).  Cysticercus  of  the  eye  deserves  special  mention; 
in  rare  cases  the  cysticercus  has  been  met  with  in  the  subcutaneous 
cellular  tissue  of  the  eyelid,  once  in  the  muscle  bundles  of  the 
musculus  orbicularis.  Subconjunctival  cysts  are  found  chiefly  in 
youthful  individuals.  Their  position  is  most  varied,  generally  in  the 
neighbourhood  of  the  inner  angle  of  the  eye.  Dilated  vessels  pass 
right  over  the  cysts,  which  are  generally  movable,  together  with  the  base 
they  rest  upon,  producing  a  spherical  protrusion.  The  head  of  the 
worm  can  sometimes  be  seen  shining  through  as  a  whitish  speck. 
The  only  symptoms  are  those  of  a  slight  irritation  of  the  connective 
tissue  and  some  difficulty  in  closing  the  lid  ;  larger  cysts  dislocate  the 
globe.  The  diagnosis  has  the  rapid  growth  of  the  cystic  tumour  to 
support  it ;  there  is  the  possibility  of  its  being  mistaken  for  a  foreign 
body  (Kaldrovils^).  After  division  of  the  connective  tissue  capsule 
extraction  is  easily  performed.  It  is  most  rare  for  the  cysticercus  to 
occur  in  the  orbit.  Suppuration  of  the  cyst  may  have  serious  con- 
sequences for  the  eye.  It  is  only  exceptionally  that  the  cysticerci 
gain  access  to  the  anterior  chamber  of  the  eye. 

Subretinal  cysticerci  or  those  localized  in  the  vitreous  are  more 
frequent.  Upon  examination  with  the  ophthalmoscope  there  is  seen 
in  the  vitreous  a  bluish  bladder  with  a  smooth  surface.  The  head  is 
seen  as  a  white  patch,  and  the  circle  of  hooks  and  the  suckers 
also  come  into  view,  also  the  frequent  movements  which  the  head 
and  neck  make  in  the  vitreous.  Operation  generally  yields  good 
results  ;  in  rare  instances  the  globe  is  atrophied  and  must  be 
enucleated. 

Formerly  cysticerci  in  the  brain  were  met  with  in  fair  frequency, 
but  the  number  of  such  cases  has  generally  decreased  of  late  years  in 
a  remarkable  way,  in  correspondence  with  the  diminution  of  cysticerci, 
which  is  to  be  attributed  to  compulsory  meat  inspection.  Whilst, 
for  example,  the  post-mortem  records  of  the  Pathological  Institute  in 
Berlin  before  the  year  1875  showed  20  per  cent,  cysticerci  affecting 
the  brain,  this  number  declined  later  to  16*3  per  cent.,  and  of  late 
years  has  fallen  to  i  per  cent.  (Orth^).  Nevertheless  even  now 
cysticercus  still  plays  no  inconsiderable  part  in  the  etiology  of 
cerebral  diseases.  For  example,  in  the  clinic  of  de  Amicis  at  Naples, 
among  seven  cases  of  cysticerci  of  the  skin,  they  were  found  four 
times  also  in  the  brain  (Sipari*).  Cysticerci  may  occur  in  the  dura 
mater,  arachnoid,  pia  mater,  choroid  plexus,  the  surface  of  the  cerebral 

*  Huber,   **  Bibliographic  der  klin.  Helminthologie,"  1891,  pt.  2. 

-  Kaldrovils,  Wien.  med.   Wochenschr.^  ig02. 

2  Orth,  Berl.  med.  Ges.,  June  29,  1904.  *  Sipari,  "  Angelo  Trani  Neapel,"  1900. 


SUPPLEMENT  665 

hemisphere,  the  medullary  substance,  the  ventricles,  the  aqueduct, 
the  corpus  striatum,  corpora  quadrigemina,  the  pineal  gland,  the  pons, 
the  cerebellum,  the  olfactory  trigone,  the  bulb,  the  medulla  oblongata, 
and  the  olive.  They  are  most  frequently  found  in  the  cortical  sub- 
stance and  in  the  ventricles ;  the  frequency  of  the  latter  situation  may 
be  explained  by  the  flow  of  the  fluid  (Henneberg^).  The  severity  of 
the  symptoms  is  not  always  in  proportion  to  the  number  of  cysticerci. 
Cases  have  been  known  in  which  ten,  twenty  and  forty  cysticerci  have 
been  found  (Hagen-Thorn^),  and  yet  the  clinical  symptoms  have 
been  remarkably  slight.  On  the  other  hand,  solitary  cysts  may 
both  run  a  course  completely  without  symptoms  and  also  cause  the 
severest  symptoms  when  located  in  specially  important  parts  of 
the  brain  (crus,  pons,  central  convolutions).  In  the  case  mentioned 
by  Jacobson^  the  invasion  of  the  brain  by  cysticerci  was  immense  ; 
the  largest  cyst  was  found  in  the  cerebral  cortex.  The  chief 
symptoms  of  cysticercus  of  the  brain  substance  consist  in  the 
onset  of  cortical  epilepsy,  which  sometimes  runs  a  very  pernicious 
course,  frequently  with  psychical  disturbances,  whilst  paralyses  are 
absent.  Perhaps,  too,  the  localization  of  pain,  spontaneous  and  on 
pressure,  corresponding  with  the  points  observed  on  the  cranium, 
is  of  importance.  Cysticerci  may  also  change  their  position 
in  the  brain  ;  patients  who  had  earlier  suffered  from  epileptiform 
convulsions  later  showed  intra-ocular  cysticerci  after  the  cerebral 
symptoms  had  completely  disappeared.  Treatment  can  only  be 
surgical  ;  v.  Bergmann*  operated  in  two  cases  with  well-marked 
improvement.  Parasites  in  the  ventricles  are  especially  dangerous, 
more  especially  so  when  free  in  the  ventricles,  and  so  capable  of 
giving  rise  to  the  danger  of  sudden  closure  of  the  foramen  of  Majendie 
(Simmonds,^  Verse^).  Stern^  states  the  symptoms  of  cysticercus 
in  the  fourth  ventricle  to  be  the  following:  general  cerebral  pressure 
symptoms  (headache,  vertigo,  vomiting,  somnolence,  congested  disc 
caused  by  internal  hydrocephalus)  ;  in  addition,  there  are  symptoms 
which  point  to  disease  of  the  hind-brain — pain  and  stiffness  in  the 
neck,  vertigo  and  cerebellar  ataxy,  violent  and  persistent  vomiting, 
slowness  of  pulse  ;  and  lastly  those  rare  but  certain  symptoms  of  a 
lesion  of  the  bulb,  such  as  diabetes,  respiratory  disturbances  and 
paralysis  of  cerebral  nerves,  especially  of  the  abducens.  These  are 
far  less  marked  than  the  general  symptoms  of  cerebral  pressure.  One 
characteristic  is  the    remarkable  alternation  between   severe  general 

'  Ilenneberg,  Berl.  kliii.  Wochenschr.y  1906,  xxxii. 

■■^  Hagen-Thorn,  abstract  by  Posselt.  ^  Jacobson,  Berl.  klin.  IVochenschr.,  igo6. 

^  V.  Bergmann,  quoted  by  Fiangenheim,  loc.  cit.,  p,  470. 

•'■  Simmonds,  Milnch.  med.  Wochenschr.^  1907,  xxvii. 

*  Verse,  Milnch.  med.  IVochenschr.,  1907,  xi.         '  Stern,  Zeitschr.  f,  klin.  Med.,  Ixi. 


666  THE  ANIMAL   PARASITES   OF   MAN 

symptoms  and  periods  of  complete  sense  of  well-being ;  in  this  way 
a  functional  nervous  affection  may  be  simulated  (Jolasse^).  Brun's 
symptom  (in  the  widest  sense,  sudden  onset  of  violent  cerebral 
symptoms  upon  change  of  head-posture)  is  a  specially  characteristic 
sign  of  free  cysticercus  in  the  fourth  ventricle ;  the  disease  generally 
terminates  with  sudden  death  from  cessation  of  the  heart's  action. 
Defects  in  motor  power,  convulsions,  implication  of  other  nerves, 
are  rare  and  unessential  complications  (Hartmann^).  Carefully 
carried  out,  lumbar  puncture  may  possess  some  diagnostic  and 
therapeutic  value.  Treatment  is  purely  symptomatic,  or  eventually 
Neisser's  ventricle  puncture  may  be  considered. 

At  the  base  of  the  brain  the  cysticerci,  as  a  rule,  assume  that 
form  which  is  designated  as  C.  racemosns,  and  consists  of  rows  of 
delicate  grape- like  bladders  in  groups,  sometimes  also  markedly 
branched,  but  generally  sterile,  which  develop  in  the  meshes  of  the 
soft  meninges  and  may  envelop  the  nerves  and  vessels  of  the  base  of 
the  brain.  Such  tumours  bring  about  hydrocephalus  and  chronic 
leptomeningitis,  which  must  be  regarded  as  the  causes  of  the  clinical 
disturbances  (cysticercus  meningitis),  attacks  of  loss  of  conscious- 
ness, dementia  and  apathy,  dulness  and  confusion  and  headaches. 
In  the  case  recorded  by  Meyer^  symptoms  which  resembled  paralysis 
agitans  were  noteworthy,  and  defects  in  speech  in  the  case  recorded 
by  Dursf*  (C.  racemosus  in  the  region  of  the  left  Sylvian  fossa). 
According  to  Markwald^  C.  racemosus  of  the  fourth  ventricle  is  said 
to  represent  a  characteristic  clinical  picture  :  violent  headaches, 
attacks  of  vertigo  followed  very  soon  by  deep  coma  and  death 
in  a  few  days.  Treatment  in  Cysticercus  racemosus  is  ineffectual. 
In  the  diagnosis  of  cerebral  cysticerci  in  general  the  recognition  of 
multiple  cysticerci  in  the  skin  and  muscle  and  of  the  tapeworm  is 
of  importance.  In  cases  of  cerebral  diseases  in  which  cysticerci  may 
be  a  possible  cause,  Remmert*^  recommends  that  the  skin  of  the 
whole  body  should  be  palpated. 

Cysticercus  in  the  spinal  cord  and  in  the  vertebral  column  is 
occasionally  observed  ;  as  a  rule,  other  organs,  above  all  the  brain 
and  its  membranes,  are  simultaneously  affected.  Here,  too,  the 
cysticercus  occurs  in  two  forms— sometimes  the  cysts  are  roundish 
or  oval,  solitary  or  multiple,  and  at  other  times  Cysticercus  raceuwsus 
occurs. 


^  Jolasse,  Mibu/i.  med.  Wochenschr.,  1896. 

-  Ilartmann,  Wien.  klin.  Wochenschr.,  1902. 

^  Meyer,  Detitsch.  tried.  Wochenschr.,  1906.  *  Durst,  Lieen.  viestnik,  1902. 

^  Markwald,  Milnch.  med.  Wochenschr.^  1895.         ^  Remmert,   "Dissert.  Berlin,"  1893. 


SUi'PLEMENT  667 

Taenia  saginata. 

Occurs  in  the  small  intestine  of  man.  It  is  characteristic  of  the 
habit  of  life  of  this  parasite  that  once  it  has  become  mature  its 
proglottides  are  dropped  off  daily  in  increasing  numbers  because 
its  growth  is  extraordinarily  rapid.  The  joints  are  discharged 
generally  spontaneously  during  the  whole  day  without  a  stool.  An 
extraordinarily  unpleasant  sensation  is  produced  by  the  damp,  cool 
joints  slipping  down  into  one's  lower  garments  and  over  one's  legs 
when  walking ;  women  especially,  in  whom  the  proglottides  slip 
through  their  petticoats  on  to  their  legs,  complain  bitterly  of  this 
troublesome  symptom.  Another  unpleasant  symptom  is  superadded 
in  the  shape  of  the  proglottides  tickling  the  rectum,  and  this  excites 
irritable  people  to  the  last  degree.  Different  species  of  tapeworms  are 
not  mutually  exclusive.  B.  latns  and  T.  solium  frequently  occur  side 
by  side,  so  also  T.  sollnm  and  T.  saginata — for  instance,  in  a  butcher's 
assistant  w-e  once  expelled  twelve  T.  solium  and  one  T.  saginata  at 
the  same  time.  The  greatest  number  of  Taenia3  which  have  been 
observed  at  one  time  amounted  to  forty  T.  solium  (Kleefeld^). 
Even  though  the  cysticercus  of  T.  saginata  is  not,  as  in  the  case  of 
T.  solium,  particularly  dangerous  to  man,  a  parasite,  nevertheless, 
which  requires  so  much  nutrient  material  during  its  rapid  growth, 
and  thereby  sets  up  manifold  disturbances  in  the  general  condition 
of  health,  ought  to  be  expelled  as  rapidly  and  thoroughly  as  possible. 

Tapeworms  are  found  not  uncommonly  with  other  intestinal 
parasites,  such  as  Ascaris,  Oxyuris,  Trichocephalus  or  Ancylostoma. 
Prunac^  described  a  case  in  which  a  woman  passed  a  Taenia  through 
the  anus  while  she  vomited  a  Fasciola  hepatica. 

The  symptomatology  of  these  three  large  species  of  Cestodes^ 
Dibothriocephalus  latns,  Tcenia  solimn,  and  T.  saginata,  may  very  well 
be  summarized  together,  as,  apart  from  some  peculiarities,  the  clinical 
symptoms,  especially  so  far  as  their  localization  in  the  intestine  is 
concerned,  are  practically  the  same  for  all  three  species.  In  a  large 
number  of  cases  the  hosts  have  no  suspicion  whatever  that  they  are 
harbouring  a  tapeworm  ;  they  feel  quite  well  and  free  from  any 
disquieting  symptoms  whatever,  and  only  become  aware  of  the  fact 
that  they  are  the  carriers  of  a  tapeworm  when  the  discharge  of  the 
segments  takes  place;  on  the  other  hand,  it  is  often  difficult  to  rid 
people  of  the  idea  that  they  are  harbouring  a  Taenia  (Kiichenmeister 
calls  such  Taenia  imaginata) ;  usually  it  is  undigested  fibrous  shreds 
of  beefsteak  which  are  regarded  by  the  patients  as  proglottides  of 
taeniae. 


'  Kleefeld,  see  Seifert  loc.  cit. 

■^  Prunac,  see  Eichhorst,   **  Handb.  d.  spez.  Path.  u.  Therap.,"  ii,  p.  281. 


42 


668  THE   ANIMAL   PARASITES   OF   MAN 

In  a  large  number  of  cases,  disturbances  of  the  intestinal  tract 
set  in,  e.g.,  sense  of  pressure  in  the  abdomen,  which  sometimes 
becomes  constant  on  one  and  the  same  side,  or  sometimes 
changes,  now  at  the  umbilicus  and  again  at  the  epigastrium  ;  here 
and  there  colicky  pains  are  present.  Derangements  of  appetite  and 
digestion  are  frequently  complained  of ;  the  most  frequent  are  the 
sensations  of  morbid  hunger  or  irregular  appetite,  nausea  and 
vomiting.  Thus,  at  the  Third  Congress  of  Internal  Medicine, 
Senator  recorded  a  case  in  which  there  were  symptoms  of  nervous 
dyspepsia,  cured  after  a  successful  vermifuge.  There  is  either 
constipation  or  diarrhoea,  so  that  many  of  such  patients  are  brought 
for  treatment  with  the  diagnosis  of  "  chronic  intestinal  catarrh  "  and 
correspondingly  treated.  As  to  the  treatment  of  toxic  action  of  the 
Taeniae  when  such  arises,  see  the  special  section  on  the  subject 
(bothriocephalus  anaemia,  p.  644).  The  frequent  disturbances  of  the 
general  condition,  so-called  reflex  phenomena,  so  far  as  the  action  of 
toxic  substances  is  not  in  question,  may  be  explained  by  the  fact  of 
their  occurrence  in  specially  sensitive  individuals  who  are  affected  by 
such  phenomena.  The  proof  that  a  diseased  condition  is  produced 
by  a  tapeworm  will  be  forthcoming  with  some  degree  of  certainty 
if  the  symptoms  cease  immediately  after  the  removal  of  the  parasites. 
As  a  whole  series  of  troubles,  which  certainly  have  nothing  to  do 
with  them,  are  erroneously  ascribed  to  the  tapeworm,  as  is  frequently 
assumed,  one  will  do  well  to  be  somewhat  critical  in  this  respect. 

The  treatment  is  of  a  threefold  nature  :  prophylactic,  symptomatic 
and  radical. 

Under  any  circumstances,  the  best  prophylaxis  is  that  which 
consists  in  only  eating  the  flesh  of  those  animals  in  which  any  of  the 
three  larval  forms  occur  (pig,  cattle,  salmon,  pike,  burbot,  etc.) 
so  prepared  that  the  larval  forms  have  been  destroyed  and  the  food 
thus  rendered  innocuous.  For  domestic  and  public  use  the  rule 
prescribed  by  Kiichenmeister  is  under  all  circumstances  most  easily 
understood,  namely  to  roast  or  boil  till  the  flesh  appear  greyish- 
white  and  sufficiently  done  by  reason  of  the  coagulation  of  the 
albumen  and  decolorization  of  the  blood.  The  general  prophylaxis 
simply  concerns  the  tapeworm  carriers  trying  to  limit  as  far  as 
possible  the  further  extension  of  the  parasites  in  the  animal  world 
by  carefully  rendering  the  expelled  segments  and  worms  harmless 
(pouring  sulphuric  acid  over  the  faeces  and  burning  the  worms)  and 
also  by  strictly  adhering  to  official  regulations.  The  official  system 
of  meat  inspection  in  this  respect  has  been  of  immense  service,  and 
much  can  still  be  done  by  means  of  thorough  official  control  over 
cleanliness    in    abattoirs    and    butchers'    shops.     Galli-Valerio^   very 

'  Galli-Valerio,  Therap.  Monatsh. ,  1900. 


SUPPLEMENT  669 

rightly  desires  the  aboHtion  of  the  custom  of  manuring  fruit-plants 
such  as  strawberries,  vegetables  and  salad  with  the  contents  of  privies, 
and  would  extend  the  use  of  privies  in  the  country. 

Symptomatic  treatment  consists,  in  the  case  of  those  Taeniae  which 
resist  radical  attempts  at  expulsion,  of  repeated  use  of  drugs  injurious 
to  the  worm  as  soon  as  ever  new  proglottides  are  formed,  or  in 
special  cases,  as  in  the  case  of  persons  weakened  by  diseases  or 
operations,  or  frail  old  people,  or  patients  with  severe  heart  failure, 
gastric  or  intestinal  carcinoma,  or  in  pregnancy,  in  effecting  the 
expulsion  of  a  large  chain  of  proglottides  by  the  mildest  measures 
possible. 

Radical  treatment  of  the  Taenia  is  not  always  equally  easy  in  all 
three  species,  even  when  the  means  used  are  the  same ;  the  easiest  to 
expel  is  T.  solituHy  then  D.  latus,  and  the  most  difficult  T.  saginata. 
That  as  yet  no  certain  cure  exists  for  Cestodes  is  clear  from  the  large 
number  of  drugs  recommended  from  time  to  time,  and  the  increase 
of  bungling  treatment  in  this  respect ;  in  addition,  there  is  no  depart- 
ment in  which  there  is  so  much  quackery  as  in  vermifuges.  The 
treatment  proper  should  always  be  preceded  by  thorough  preparatory 
treatment,  the  purpose  of  which  is  to  render  the  gut  as  empty  as 
possible  once  for  all,  and  on  the  other  hand  to  put  the  worms  them- 
selves into  a  diseased  condition.  How  far  the  host  himself  has  been 
made  ill  by  such  preliminary  cures  (herring,  pickle,  garlic,  onions, 
preserved  strawberries),  many  a  person  who  has  had  to  do  with  such 
things  can  recount.  In  the  opinion  of  Fischer^  strict  preparatory 
treatment  appears  to  favour  the  development  of  toxic  substances,  or 
else  it  disposes  to  vomiting ;  as  a  rule  it  causes  the  patient  far  more 
discomfort  than  the  treatment  itself.  In  recent  times  far  less  weight 
is  attached  to  these  preparatory  treatments  than  to  carefully  prepared 
and  correctly  dosed  drugs  ;  the  preparation  is  generally  limited  to 
relieving  the  intestine  in  a  simple  way,  the  day  before  the  treatment, 
of  the  densest  faecal  masses,  by  a  simple  aperient  or  water  enema. 

We  recommend  the  following,  which  has  always  proved  itself  to 
be  the  best  and  simplest  remedy  against  T.  saginata.  The  pati-crnt 
takes  early  in  the  evening  before  the  treatment  nothing  but  a  plate  of 
soup  or  a  glass  of  milk,  and  then  takes  a  laxative  (electuar.  lenit  or 
infus.  sennae  compos,  or  an  enema),  so  that  later  in  the  evening  one 
to  two  stools  are  passed.  In  this  connection  we  fail  to  agree  with 
Grawitz^  and  Boas,^  who  consider  that  at  least  preliminary  evacua- 
tion   of  the    intestines    can  be    dispensed    with.     On   the   following 


'  Fischer,  Stockholm,  Nordin  and  Josephson,  1904. 
'^  Grawitz,  Munch,  med.  Wochenschr.^  1899. 
^  Boas,  Dtutsch.  med.  Wochenschr.^  i839> 


670  THE   ANIMAL   PARASITES   OF   MAN 

morning  the  patient  should  take  a  cup  of  black  coffee  or  tea  without 
anything  else,  and  half  an  hour  later  the  vermifuge. 

The  best  drug  is  extract,  filicis  maris  aether.,  which  also  forms  the 
main  constituent  of  most  of  the  secret  remedies  recommended  for 
tapeworms.  Earlier  mishaps  with  this  preparation  had  their  origin 
principally  in  insufficient  dosage.  Also,  in  addition  to  correct  dosage, 
extract,  filic.  maris  needs  very  careful  preparation  if  satisfactory  results 
are  to  be  attained.  If  preparations  wnth  the  trade  mark  ''  Helfenberg  " 
or  "Wohnar"  are  not  used,  but  the  male  fern  extract  has  been 
prepared  by  a  chemist,  one  must  make  certain  that  the  roots  of  the 
Aspidium  filix-mas  have  been  collected  in  May  or  October,  and  only 
green  sappy  specimens  selected,  and  that  the  attached  paleae  have 
been  separated,  that  they  have  been  broken  up  small  and  ether 
poured  over  them  with  a  little  spirits  of  wine  while  quite  fresh.  The 
w^hole  mass  is  to  be  kept  in  a  cool  place,  but  not  too  closely  covered. 
If  at  any  time  a  certain  quantity  is  to  be  used,  it  is  taken  out,  the 
ether  carefully  distilled  in  a  retort  till  the  extract  has  a  suitable  fluid 
consistency.  Fischer  attaches  great  importance  to  the  direction  in 
the  Pharmacopoeia  being  exactly  follow^ed,  to  the  effect  that  the 
extract  is  to  be  carefully  stirred  before  prescribing,  as  the  active  sub- 
stances undergo  partial  crystallization  if  kept  for  any  length  of  time 
and  sink  to  the  bottom,  so  that  the  preparation  has  a  different  strength 
and  toxicity  in  different  layers.  Of  this  extract  10  to  12  to  15  grm.  are 
to  be  taken  in  gelatine  capsules  within  half  an  hour.  We  consider  it 
unjustifiable  to  give  greater  doses  than  15  grm.  to  adults,  as  many 
cases  are  known  in  which  to  some  extent  severe  toxic  symptoms  have 
followed,  such  as  headache,  sensation  of  giddiness,  dyspnoea  and 
cyanosis,  yellow^  vision  (xanthopsia),  delirium,  stupor,  the  most  severe 
cramps  in  the  extremities,  rapidly  fatal  trismus  and  tetanus.  The 
most  serious  are  defects  of  vision  of  various  kinds,  which  may  end 
in  amblyopia  and  amaurosis,  with  permanent  blindness.  A  complete 
collection  of  toxicological  literature  up  to  the  year  1903  is  to  be 
found  in  Marx's^  Dissertation.  Since  that  time  further  instances 
of  such  intoxications  have  been  made  known.  NageP  observed 
them  only  in  severe  cases.  O.  Meyer^  lays  special  stress  on  the  bad 
prognosis  of  the  disturbances  of  vision  evoked  by  poisoning  \vith 
extract,  filicis  maris.  Studt^  has  seen  two  cases  of  optic  neuritis,  one 
with  circumscribed,  the  other  with  diffuse  retinal  oedema.  Uhthoff'' 
has  only  seen  one  case  ;  in  that  reported  by  Noiszewski^  the  toxic 
retinitis  w^as  cured;   in  ViereckV  case  bilateral  concentric  limitation 


'  Marx,  ••  Diss.  WUrzburg,"  1903,  2  Nagel,  Deutsch.  med.  Wochenschr.,  1903. 

^  Meyer,  O.,  i5<?r/.  klin.  Wochenschr.,  1905.  *  Studt,  ibid.,  1905. 

•"'  Uhthoff,  ibia.,  1905.  «  Noiszewski,  "  Postepokuhs',"  190''). 

'  Viereck,  Arch.  f.  Schiffs- ti.  Tropen-Hyg.,  1906. 


SUPPLEMENT  67 1 

of  the  field  of  vision  followed  three  days  after  taking  8*o  grm.  extract, 
filicis  maris.  Stuelp^  attributes  the  amaurosis  occurring  after  taking 
fihx  mas  to  a  toxic  action  on  the  muscularis  of  the  central  retinal 
artery  ;  there  followed  paralysis  of  the  vessel,  vascular  engorgement, 
and  thereby  nutritional  defects  of  the  nervous  elements  followed. 
In  children  one  has  to  diminish  the  dose  correspondingly,  as  with 
them,  still  more  so  than  with  adults,  severe  disturbances  arise.  Huber^ 
claims  that  this  drug  should  not  be  given  to  children  indiscriminately. 
The  view  is  frequently  expressed  that  a  combination  of  extractum 
filicis  maris  with  fatty  oils  in  which  the  active  constituents  are  soluble 
favours  intoxication.  Marx^  also  argues  from  this  standpoint  and 
assumes  that  the  ideal  preparation,  free  from  objection,  would  be  got 
if  from  filix-mas  extract  a  preparation  free  from  fatty  oils  could  be 
made,  and  he  considers  it  advisable  to  limit  the  use  of  castor  oil  as  an 
aperient  before  and  after  taking  the  "  cure,"  and  to  prescribe  instead 
a  saline  laxative,  such  as  Epsom  salts  or  Glauber's  salts.  Sonnen- 
schein^  also  advises  against  the  simultaneous  exhibition  of  extractum 
filicis  maris  with  oleum  ricini,  as  is  the  case  with  Helfenberg's 
capsules,  and  Boas^  is  likewise  anxious  that  ol.  ricini  should  be 
avoided.  Lenhartz^  appears  to  consider  the  warning  against  the 
simultaneous  combination  of  the  extract  with  fats  or  ethereal  oils,  and 
especially  against  the  employment  of  castor  oil  as  an  after-treatment, 
as  without  justification,  and  we,  too,  in  the  course  of  our  many 
filix  treatments,  have  never  yet  witnessed  any  unfavourable  effect  from 
the  use  of  castor  oil  in  the  after-treatment.  The  surest  way  of 
obviating  the  toxic  effects  of  extractum  filicis  is  to  give  a  laxative 
(ol.  ricini)  as  soon  as  the  extract  has  left  the  stomach,  say,  about  half 
an  hour,  so  that  it  need  not  stay  longer  than  necessary  in  the  gut  and 
become  absorbed.  Perhaps  in  most  cases  of  poisoning,  transgressions 
against  this  rule  have  been  the  cause  of  the  toxic  action.  The  nausea 
that  sets  in  the  day  after  taking  the  drug  and  the  inclination  to  vomit 
are  best  resisted  by  giving  iced  coffee,  iced  tea,  iced  pills,  peppermint 
tea,  cognac,  one  to  two  wafer  powders  of  menthol  and  sacch.  lactis 
aa  0'2  grm.  (Apolant*^)  half  an  hour  before  the  drug  is  taken. 
Fischer*^  considers  that  lying  still  in  the  horizontal  position  is  the 
best  remedy.  Boas^  recommends  the  injection  of  the  drug  into  the 
stomachs  of  patients  who  tolerate  extractum  filicis  badly,  in  the  form  of 
a  thin  emulsion  (with  gi.  arab.).     In  the  case  of  children  the  extract 


'   Stuelp,  Arch.  f.  Au^enheilk.,  1906,  li. 

'^  YlnhQXy  Miinch.  med.  Wochenschr.,  1903.  ^  Marx,  loc.  cit. 

4  Sonnenschein,  Milnch.  med.  Wochenschr.,  1903-  ^  Boas,  loc.  cit. 

^  Lenhartz,  loc.  cit,  "'  Apolant,  Deutsch.  med.  Wochenschr..^  1905,  xHv. 

^  Fischer,  loc.  cit.  ^  Boas,  l<yt.  cit. 


6/2  THE   ANIMAL   PARASITES   OF   MAN 

is  prescribed  with  honey  as  an  electuary.  The  method  recommended 
by  Fowler'  is  without  doubt  too  detailed  ;  he  prescribes  before  the 
treatment  two  to  three  to  four  days'  rest  in  bed ;  special  diet,  tablets  of 
cascara  sagrada  three  times  daily,  on  the  fourth  day  senna  infusion, 
and  then  to  give  the  extractum  lilicis  maris  in  capsules  in  four  doses, 
to  be  taken  every  quarter  of  an  hour. 

Under  Jaquet's^  direction,  Kraft  has  prepared  an  amorphous  acid 
from  the  fern  root  extract  which  is  designated  filmaron.  As  a  vermi- 
fuge the  drug  is  prescribed  for  children  of  2  to  5  years  of  age  in 
doses  up  to  o'2  to  0*3  grm.,  for  children  of  from  8  to  12  years  in  doses 
up  too'5  too7grm.,and  for  adults  up  to  07  to  I'o  grm.,  so  as  to  expel 
the  parasites.  Bodenstein^  gives  the  tilmaron  oil  introduced  into 
commerce  by  the  firm  of  Boehringer  (one  part  filmaron  and  nine  parts 
castor  oil)  in  still  greater  dosage,  either  fasting  or,  in  the  case  of 
sensitive  patients,  one  hour  after  a  cup  of  tea ;  he  gives  peppermint 
tablets  against  possible  nausea.  Brieger^  tested  the  preparation  in 
twenty-three  cases  ;  in  twenty-one  of  these  he  prescribed  it  as  an  ether- 
castor  oil  mixture,  and  in  two  as  capsules.  The  action  always  took 
effect  in  from  two  to  five  hours,  and  only  in  three  cases  were 
unpleasant  after-effects  in  the  shape  of  colic  observed  ;  in  sixteen 
cases  the  result  was  positive,  in  seven  negative. 

The  attempts  made  by  Goldman n^  to  prepare  from  the  bark  of 
Miisemia  ahyssinka,  a  plant  of  the  order  Myrsijiacece,  indigenous  to 
Persia,  the  active  substance,  namely  sebirol,  have  shown  that  when 
this  is  given  alone  it  certainly  acts  as  a  vermicide,  but  not  as  a 
vermifuge  ;  on  the  other  hand,  the  results  of  a  combination  of  sebirol 
with  thymol  and  salicylates  were  surprisingly  good  ;  this  mixture  has 
been  introduced  into  commerce  as  taeniol,  in  the  shape  of  pastilles 
prepared  with  chocolate  for  children.  The  method  of  giving  taeniol 
is  as  follows  :  On  the  day  before  the  administration  a  light  diet  and 
thorough  purging  wath  calomel  are  ordered  ;  and  then  on  the  day  of 
the  treatment  itself,  after  a  breakfast  consisting  of  a  cup  of  tea,  in  the 
case  or  adults,  thirteen  to  fifteen  taeniol  pastilles  are  taken  in  some  red 
wine  at  intervals  of  ten  minutes  respectively.  In  the  middle  of  this- 
treatment  an  interval  of  some  hours  is  interposed.  After  the  pastilles 
have  been  taken  a  calomel  purge  is  again  given.  The  results  obtained 
by  Liermberger^  are  sufficiently  encouraging  to  be  put  to  further  test. 

Fischer''  has  tested  in  some  of  his  cases  extracts  of  some  new 
species  of  fern  root ;  he  employed  the  extract  from  the  rhizomes  of 


1  Fowler,  B'//.  ^r^,{.  Jonrn.,  1906.  -  Jaquet,  Therap.  Monatsh.,  1904. 

»  Bodenstein,  IVieu.  vied.  Presie,  1906.  ^  Brieger,  "Therap.  d.  Gegenwart.,"  1905. 


^  Goldmann,  Wien.kiin.  Wocheuschr.,  1905. 

^  Liermberger,  BerLklin.   llocliemchr.,  ito5.    '    Fischer,  loc.  cit. 


SUPPLEMENT  673 

Aspidium  spimilosiim  and  ^.  dilatatum,  two  fern  roots  indigenous  to 
Sweden,  and  obtained  remarkable  results  (doses  of  4  grm.).  Lauren^ 
had  previously  recorded  similar  results,  and  recently  Friedjung,^ 
using  extr.  aspid.  spinulos. 

Cortex  radicis  granati  as  fresh  bark  is  a  very  good  drug,  and  is 
usually  given  as  a  decoction  :  iSo'O  bark  to  i,ooo*o  water,  boiled  for 
forty  hours  to  240-0,  and  a  small  cupful  to  be  given  every  half  an  hour  ; 
colic,  vomiting  and  diarrhoea,  are,  however,  easily  induced.  The  chief 
constituent  of  the  granate  root,  pelletierinum,  possesses  vermicidal 
properties,  and  is  much  recommended,  especially  in  France.  Sequelae 
easily  arise  (vertigo,  hazy  vision,  malaise,  vomiting,  quickened  heart's 
action,  muscular  tremors,  cramps  in  the  calves),  especially  in  delicate 
persons  and  children,  so  that  one  should  refrain  from  giving  it  to  the 
latter  especially  (Drivon^).  Sometimes,  judging  by  the  experience  of 
Sobotta^  and  Boas,-'^  the  action  is  problematical.  Where  it  is  desired 
to  employ  it  in  the  case  of  adults,  the  following  is  prescribed  :  pellet, 
sulfur.  0-3  to  o*4  grm.,  acid,  tannic.  0*5  grm.,  sir.  rub.  jd.  30*0  grm.,  to 
be  taken  at  one  time,  and  a  quarter  to  half  an  hour  after  a  purgative 
(senna  infusion).  In  the  case  of  children  it  is  better  to  employ 
semina  cucurbitae  maximae  instead  of  extractum  filicis  maris.  Sixty 
to  100  pumpkin  seeds  are  pounded  up  with  sugar,  which  yield  a 
pleasant-tasting  electuary,  and  which  are  taken  all  at  once  ;  half  an 
hour  afterwards  a  laxative  is  taken  (Storch,^  Pick^).  Jungklauss's 
preparation  is  nothing  else  than  a  pumpkin  extract ;  its  action  is 
favourable;  it  is,  however,  too  expensive  (Ritter^).  Flores  kousso 
up  to  15  to  20  grm.  in  compressed  form  or  in  sugar  or  honey  in  the 
form  of  electuaries  (children  2-0  to  10*0  grm.  according  to  age)  is  not  to 
be  relied  upon ;  kussin,  prepared  from  kousso  flowers  (Bedall,  Munich), 
is  not  a  pure  body ;  when  taken  it  is  divided  into  four  parts  up  to 
ro  to  2'o  grm.  with  elaeosaccharum  menthae,  at  half-hourly  intervals  ; 
it  is  said  to  be  less  unpleasant  than  treatment  with  flores  kousso  (Lieb- 
reich  and  Langgard^).  Kosinum  crystallisatum  (dose  1*5  to  2'o  grm.) 
is  prepared  by  the  firm  of  Merck.  Kamala  is  the  least  potent  of  the 
tapeworm  drugs  in  use,  and  is  principally  to  be  recommended  in  the 
treatment  of  children:  1*5  to  3*0  grm.  in  electuaries.  According  to 
Leichtenstern^^   and  White  ^'  chloroform,  even  in  toxic  doses,  cannot 


'  Lauren,  Therap.  Monatsh.^  1899. 

2  Friedjung,  Ges.  f.  innere  Med..,  Wien,  March  8,  1906. 

•^  Drivon,  Lyon  med.,  1902.  ■*  Sobotta,  loc.  cit.  *  Boas,  loc.  cit. 

^  Storch,  see  Lenhartz,  loc.  cit. 

■^  Pick,  Ges.  f.  innere  Med.,  Wien,  March  8,  1906. 

^  Ritter,  Frag.  med.  IVochenschr.,  1904,  v. 

^  Liebreich  and  Langgard,  '*  Kompendium  der  Arzneiverordnung, "  1907. 
'°  Leichtenstern,  *'  Therap.  der  Gegenwart.,"  1899. 
"  White,  Scot.  Med.  and  Surg.  Journ.,  1900. 


674  THE   ANIMAL  PARASITES   OF   MAN 

do  any  harm  to  the  tapeworm,  nevertheless  it  has  been  recently 
recommended  by  Carratu^  ;  chloroform  6'o,  sirup.  6o'o,  one  tea- 
spoonful  to  be  taken  every  hour  (fasting).  Salol  is  recommended  by 
Galli-Valerio^  as  an  absolutely  harmless  tapeworm  drug ;  thymotal 
(a  derivative  of  thymol)  by  Pool,^  3  grm.  to  be  given  up  to  three  to 
four  times  on  four  consecutive  days. 

The  drug  well  known  long  ago,  cuprum  oxyd.  nigr.,  has  been 
recently  brought  into  fresh  notice  by  Dorr.*  It  is  also  the  chief  con- 
stituent of  the  tapeworm  drug  introduced  into  commerce  by  the  firm 
of  Dehlsen  (Itzehoe)  (Koch^).  The  coconut  is  absolutely  ineffectual, 
also  naphthalin,  croton-chloral,  ether,  gallanol,  strontium  lactate, 
glycerine  and  bromide  of  potash. 

Where  possible  one  should  endeavour  to  discover  the  head  or  the 
heads  of  the  tapeworm  in  the  stools,  so  as  to  make  certain  whether 
the  treatment  has  been  successful ;  this  search  is  best  carried  out  by 
immediately  and  carefully  pouring  water  over  the  total  quantity  of 
evacuations  collected  in  the  night  stool,  without  stirring  them  up,  till 
only  the  tapeworm  is  found  lying  at  the  bottom  of  the  vessel. 


NEMATODES. 

Strongyloides  stercoralis. 

The  pathological  significance  of  this  intestinal  parasite  is  not  yet 
fully  demonstrated.  In  Seifert's^  observation,  on  what  Leichtenstern' 
called  the  celebrated  Wlirzburg  case,  the  patient  had  suffered  many 
times  from  attacks  of  blood-stained  diarrhoea  with  tenesmus,  as  in 
ZinnVcase  of  a  three  year  old  boy  who  had  bloody  purulent  diarrhoea. 
Schliiter^  speaks  of  a  haemorrhagic  enteritis  produced  by  Strongyloides. 
In  other  cases  besides  diarrhoea  (either  with  or  without  blood)  there 
were  noted  :  pains  in  the  body  (Schluter),  tenderness  of  the  abdomen, 
loss  of  appetite,  gastric  troubles  of  a  general  kind,  headache,  giddi- 
ness, fainting  attacks,  anaemia  (Silvestri,^^  Valdes,^^  and  Trappe^^),  so 
that  even  if  in  isolated  cases  (Fulleborn^^)  symptoms  are  absent,  some 
significance  cannot  be  denied  these  parasites  as  a  matter  of  course 


^  Carratu,  Giorn.  med.  del  regio  eserc,  1903. 

2  Galli- Valeric,  Therap.  Monatsh.,  1900. 

'^  Foo],  Med.  Woche,  1901.  ^  Dorr,  "Therap.  der  Gegenwart,"  1901. 

^  Koch,  Med.  Klinik,  T907. 

^  Seifert,  "  Sitzungsberichte  der  phys.-med.  Ges.  in  Wlirzburg,"  1883. 

■^  Leichtenstern,  Arbeiten  aus  d.  kaiser  I.  Gesundheitsamte^  1905,  xxii. 

"  Zinn,  Berl.  klin.  IVochenschr.,  1900,  xlix. 

"  Schluter,  "  Diss.  Kiel,"  1905.  'o  Silvestri,  see  Schluter  loc.  cit.         "  Valdes,  ibid. 

'^  Trappe,  Deutsch.  rued.  IVochenschr.^  1907. 

'3  Fulleborn,  Biol.  Abt.  d.  arztl.-Vereins  in  Haviburg,  October  14,  1902. 


SUPPLEMENT  675 

(Bruns/  Leichtenstern^).  According  to  Kmiow,'  in  Siberia  there  is  a 
form  of  sporadic  bloody  diarrhoea  which  has  its  origin  in  the  presence 
of  Strongyloides  stcrcoralis.  The  parasite  does  not  Hve  only  in  the 
intestinal  lumen,  but  also  in  the  intestinal  wall,  where  it  causes 
abscesses,  fistulae  and  effusions  of  blood. 

Diagnosis  is  easily  made  by  the  detection  of  the  actively  moving 
larvae  in  the  stools. 

Treatment  is  rather  difficult,  as  it  is  not  always  successful  in 
getting  rid  of  the  parasites.  Authors  differ  as  to  the  effectiveness  of 
extr.  fil.  maris.  Goldman n^  still  considers  this  preparation  as  the  most 
effective  ;  he  recommends  preliminary  treatment  with  calomel  0*2  grm. 
and  tuber,  jalapae  0-5  grm.  a  day  before  the  special  treatment,  which 
consists  of  gelatine  capsules  of  15-0  grm.  extr.  fil.  maris  (to  be  taken 
in  the  course  of  four  hours) ;  afterwards  rectified  oil  of  turpentine 
in  gelatine  capsules.  The  thymol  treatment  {vide  Ancylostomiasis, 
p.  682),  thymol  alone  or  in  combination  with  calomel  (Schluter,^Valdes,^ 
Soussino,^  Goldmann^),  has  often  caused  diminution  of  the  number 
of  larvae,  but  also  often  remains  resultless.  Teissier^  maintains  that  by 
degrees  he  procured  complete  cure  by  the  administration  of  mercury 
in  the  form  of  blue  pill.  In  our  case  neither  thymol  nor  calomel, 
santonin,  extr.  fil.  maris,  decoct,  rad.  granat.,  had  any  result  whatever. 
Davaine''^  believes  he  attained  decrease  and  final  disappearance  of 
the  larvae  by  protracted  milk-cure.  Santonin,  tannalbin  and  other 
preparations  seem  ineffectual.  Tannin  enemata  (Mildner^^),  high 
injections  w^ith  starch  enemata  (Schliiter^^),  may  alleviate  in  persistent 
diarrhoea.  Travellers  who  are  visiting  regions  the  native  home  of 
Strongyloides  must  exercise  the  most  extreme  care  and  scrupulous 
cleanliness,  and  these  are  also  necessary  in  patients  already  suffering 
from  Strongyloides,  to  prevent  auto-reinfection  (Trappe^^). 

Dracunculus  medinensis  (Dracontiasis). 

The  guinea  worm  develops  in  the  dermis  of  human  beings  with- 
out any  symptoms  ;  only  when  it  is  completely  grown  does  it  form 
boil-like,  extremely  painful  abscesses,  in  the  greater  majority  of  cases 
in  the  legs,  in  the  region  of  the  ankle,  and  is  accompanied  by  general 
disturbance  and  a  feeling  of  heaviness,  dragging  and  pricking  of  the 


'  Bruns,  Miinch.  med.  Wochenschr.^  1907,  xix. 

-'  Leichtenstern,  Deutsch.  med.  Wochenschr. ,  1898. 

^  Kurlow,  Centralbl.  f.  Bakt.,iqo2,         ^  Goldmann,  Deutsch.  Aerzte-Zeitg.^  1903. 

•^  Schliiter,  "Diss.  Kiel,"  1905.  ^  Valdes,  loc.  cit.   '^  Soussino,  see  SchliUer  loc.  cit. 

"  Goldmann,  loc.  cit.  "  Teissier,  Arch.  d.  Med.  exp.^  1895. 

'0  Davaine,  see  Seifert,  Deutsch.  med.  Zeitg.,  1885. 
1'   Mildner,  Berl.  med.   Ges.,  July  24,  1907. 
'2  Schluter,  loc.  cit.  '^  Trappe,  loc.  cit. 


676  THE   ANIMAL   PARASITES   OF   MAN 

affected  part ;  it  occurs  more  rarely  in  the  arms,  certain  parts  of  the 
back,  the  head,  neck,  scrotum  and  penis  ;  in  a  superficial  position 
the  worm  can  occasionally  be  felt  through  the  skin.  In  most  cases 
there  is  only  one  worm  and  one  abscess,  but  here  and  there  one  finds 
patients  with  three,  four  or  even  up  to  eight  worms,  and  very  excep- 
tionally still  more,  as  in  the  cases  described  by  Poupee-Desportes^ 
(fifty  worms)  and  by  Harington^  (seventeen  worms). 

Diagnosis  offers  no  difficulty  when  the  worms  are  presenting  or 
can  be  felt  under  the  skin. 

The  inhabitants  of  the  native  home  of  the  guinea  worm,  as  a  rule, 
quietly  wait  till  it  has  got  so  far  out  that  it  can  be  conveniently 
grasped ;  it  is  then  bound  round  with  thread  and  fastened  between 
the  tips  of  a  split  piece  of  wood  and  slowly  wound  out.  In  ten  to 
twelve  days  it  can  be  wound  out  in  this  way.  Emily^  makes  injec- 
tions of  a  I  in  1,000  solution  of  sublimate  either  in  the  neighbour- 
hood of  the  worm  or  directly  into  its  body.  Mense^  managed  to 
remove  the  worm  in  one  sitting  by  laying  a  wad  of  cotton  wool 
soaked  in  chloroform  on  the  exposed  portion,  thus  stupefying  it. 
Our  therapeutic  observations  (Frangenheim^)  favour  the  free  laying 
open  of  the  existing  abscess  and  the  consequent  complete  extraction 
of  the  worm. 

Prophylaxis  depends  on  care  in  the  use  of  water  in  the  guinea 
worm  countries,  especially  dangerous  being  permanent  waters  infested 
by  Cyclops  sp. 

Filaria  bancrofti. 

The  parasitism  of  this  filaria  leads  to  the  formation  of  lymphangitis, 
elephantiasis,  chyluria,  orchitis,  chylocele,  abscesses,  lymphatic  varices, 
perhaps  also  to  chylous  ascites  and  chylous  diarrhoea. 

Lymphangitis  usually  attacks  the  extremities,  beginning  generally 
with  a  rigor  and  swelling  of  the  lymphatic  vessels  with  adjoining 
lymph  glands.  The  lymphatics  become  hard,  knotty  and  extremely 
painful,  the  overlying  skin  red  and  swollen  in  longitudinal  lines 
(Looss),  high  fever  sets  in  with,  to  some  extent,  severe  general  dis- 
turbance. After  some  days  the  attack  subsides,  the  swelling  then 
partially  disappears,  but  not  completely,  and  often  abscesses  develop 
in  consequence  of  the  lymphangitis.  Children,  as  a  rule,  suffer  from 
such  lymphangitic  attacks  (Finucane^). 

Diagnosis  is  not  easy,  for  many  other  causes  frequently  produce 
lymphangitis. 

'  Poupee-Desportes,  see  Looss,   "  Handb.  d.  Tropenkrankh.,"  1905,  i. 

2  Harington,  Brit.  Med.  Journ.^  1906.         ^  Emily,  see  Looss  loc.  cit.         *  Mense,  I'di'd. 

^  Frangenheim,  Volkmann's  Samml.  klin.   Vortriige,  424. 

^  Finucane,  Lancet,  1907. 


SUPPLEMENT  677 

Treatment  consists  in  rest,  raising  the  affected  limb,  applications 
of  vinegar  and  alum  or  liquor  plumbi,  in  some  cases  incisions  into  the 
swollen  part  under  antiseptic  precautions. 

Elephantiasis  (Arabum)  is  usually  situated  in  the  lower  extremities, 
in  men  in  the  scrotum  and  penis,  in  women  in  the  labium  pudendi, 
mons  veneris,  and  the  mammae  ;  more  rarely  it  attacks  the  upper 
extremities  or,  indeed,  the  head.  The  disease  develops  during  repeated 
attacks,  which  occur  at  irregular  intervals  of  weeks,  months  or  years, 
of  fever  accompanied  by  symptoms  of  lymphangitis  and  erysipelas 
{elephantoid  fever),  and  especially  as  the  result  of  different  accidental 
occurrences  such  as  chills,  bodily  exertions,  external  irritation.  The 
extremities  become  shapeless,  heavy  cylinders,  the  scrotum  occasionally 
a  colossal  tumour,  the  female  genitalia  and  the  mammae  smaller  or 
larger  tumours;  the  penis  often  shares  in  the  general  thickening,  the 
inguinal  glands  form  large  hard  prominent  masses,  and  enormous 
deformity  is  caused.  The  cause  is  more  often  seen  in  men  than 
women,  rarely  in  children  over  10,  never  in  younger  children. 

Treatment  of  elephantiasis  of  the  extremities  consists  in  raising  the 
affected  part,  massage,  bandaging,  vapour  baths  ;  the  large  elephantoid 
tumours  of  the  genitalia  and  mammae  can  only  be  treated  by  operative 
removal. 

Chyluria  (haemato-chyluria),  as  a  rule,  begins  by  a  series  of  attacks 
and  often  ceases  for  weeks  or  months,  the  attacks  being  accompanied 
by  fever,  pain  in  the  back  and  lumbar  region,  about  the  kidneys  and 
in  the  perinaeum.  The  attacks  are  separated  b}^  intervals  of  months' 
or  even  years'  duration,  a  continuous  chyluria  being  quite  rare.  The 
disease  may  last  many  years  without  the  constitution  being  markedly 
weakened,  but  in  other  cases  anaemia  and  debility  ensue  and  result 
in  death  from  marasmus.  In  chyluria  the  urine  becomes  completely 
opaque  like  milk ;  but  sometimes,  from  the  presence  of  bloo.d,  is  of  a 
peach-like  redness  :  the  sediment  contains  clotted  blood,  and  micro- 
scopically one  finds  fine  dust-like  fat  granules  and  red  cells  and 
leucocytes,  and  usually,  but  not  always,  filaria  larvae.  Sclerodermia 
may  possibly  be  caused  by  Filaria  (Bancroft^). 

Treatment  consists  in  administration  of  ol.  santali,  methylene  blue 
(o'i2  grm.  dose  several  times  daily),  ichthyol  (in  pills  from  0*5  to 
1*5  grm.  per  day),  ol.  terebinthinae  (0*5  to  1*5  gr.  per  day),  thymol 
(Ziemann^  had  no  result  from  either  thymol  or  methylene  blue), 
together  with  absolute  rest  in  bed,  diminution  of  all  fatty  nourish- 
ment and  administration  of  light  purgatives. 

Orchitis  is  in  acute  attacks  a  relatively  frequent  symptom  in 
the  East  ;  the  chylocele  is   rarely  marked  ;  the  fluid  usually  shows 


Bancroft,  Lancet,  1885.  -  Ziemann,  Detdsch.  vied.  Wochenschr.,  1905,  xi. 


678  THE   ANIMAL   PARASITES   OF   MAN 

numerous  larvae  ;  in  the  case  of  abscesses  they  are  generally  caused 
directly  by  the  adult  parasites,  as  they  have  often  been  found  in  them ; 
varices  of  the  lymphatic  vessels  are  either  superficial  or  deep  ; 
lymphorrhagia  arises  from  rupture  of  the  dilated  vessels  ;  chylous 
ascites  and  chylous  diarrhoea  may  also  be  produced  by  Filariae. 

Loa  loa. 

Loa  loa,  according  to  modern  investigations,  is  a  parasite  of  the 
subcutaneous  connective  tissue  of  man,  and  its  appearance  in  the 
conjunctiva  somewhat  accidental  ;  in  earlier  times  it  seems  to  have 
been  less  common  (Ziemann^).  A  number  of  cases  are  seen  in 
Europe  of  patients  who  have  lived  in  lilaria  regions,  and  on  return 
have  been  found  to  have  this  Nematode  in  the  subconjunctival  tissue. 
Pick,2  in  the  case  of  a  man  who  had  lived  in  the  Cameroons, 
found  the  parasites  in  active  motion  under  the  connective  tissue  of 
the  eyeball  right  over  the  cornea ;  extraction  was  easy.  Ziemann^ 
noted  three  cases  of  Loa  loa  in  the  eye  accompanied  by  temporary 
migratory  swellings  in  different  parts  of  the  body.  In  one  case, 
observed  by  Wurtz  and  Cleri^  (a  woman  from  the  French  Congo), 
Loa  loa  was  the  cause  of  intermittent  elastic  swellings  in  the 
subcutaneous  and  subconjunctival  tissue  (marked  eosinophilia).  In 
the  case  recorded  by  Pollack'*  (for  thirty  years  police  commissioner 
in  the  Cameroons)  the  worm  under  the  connective  tissue  of  the  left 
eye  by  its  snake-like  movements  caused  an  unpleasant  itching.  With 
cocaine  and  adrenalin  the  worm  can  be  made  visible,  and  by  means 
of  a  strabismus  hook  can  be  drawn  out  of  a  small  wound  in  the 
connective  tissue.  Martens^  exhibited  a  Filaria  extracted  from  the 
eyelid  under  local  anaesthesia. 

Trichuris  trichiura. 

Whilst  many  authors  consider  the  whip-worm  as  a  harmless 
parasite  of  the  large  intestine  (Leichtenstern/  Eichhorst/ 
Askanazy^),  the  number  of  severe  and  even  fatal  cases  of  diseases 
caused  by  it  (trichocephaliasis)  increase  so  much  that  the  Trichuris 
trichiura  must  be  excluded  from  the  group  of  harmless  intestinal 
parasites.     (For  disturbances  of  the  nervous  system  and  of  the  blood 

'  Ziemann,  Deutsch.  mcd.  IVochenschr.,  1905. 

-'  Pick,  ibid.  3  Ziemann,  loc.  cit. 

'  Wurtz  and  Cleri,  Arch.  Mid.  exper.,  1905,  ii. 

^  Pollack,  Berl.  ophthal.  Ges.,  May  17,  1906. 

^  Martens,  Berl.  vied.  Ges.,  July  24,  1907. 

'  Leichtenstern,  *' Handb.  d.  Therap.  v,  Pentzoldt-Stintzing." 

"  Eichhorst,  "  Handb.  d.  Spez.  Path.  u.  Therap." 

"  Askanazy,  Deutsch.  Arch.  f.  /din.  Med.,  1896. 


SUPPLEMENT  679 

[anaemia]  from  trichocephaliasis,  see  p.  650).  Infection  in  human 
beings  results  from  the  eggs  that  have  developed  outside  the  body, 
which  probably  reach  the  digestive  tract  on  the  hands  soiled  with  dirt 
or  earth,  or  possibly  through  drinking  water.  (Moosbrugger^  and 
Kahane^  mention  in  their  cases  that  the  children  had  an  absolute 
passion  for  earth-eating.)  Possibly,  too,  patients  reinfect  themselves 
anew,  as  an  intermediate  host  is  not  necessary. 

The  anterior  part  of  the  body  of  the  parasite  is  usually  fixed  in  the 
mucous  membrane,  and  according  to  Askanazy  feeds  on  the  blood 
of  its  host.  Moosbrugger,^  Schulze,^  Kahane,^  Vix,'^  Girard^  and 
Blanchard^  all  found  changes  in  the  mucous  membrane  of  the  gut, 
showing  that  the  parasites  had  been  in  the  gut  for  a  considerable 
time.  Kahane^  had  an  opportunity  of  seeing  at  the  Pasteur 
Institute  Trichocephali  with  the  anterior  part  of  the  body  penetrating 
not  only  the  mucosa  but  also  deep  into  the  muscularis  of  the  gut 
wall.  From  this  mode  of  attachment  to  the  wall  it  is  easily  under- 
stood how  Trichocephali,  especially  when  they  are  numerous  in  the 
gut,  cause  local  irritation  and  inflammatory  conditions  consisting  of 
frequent  attacks  of  diarrhoea,  sometimes  twenty  times  a  day,  lasting 
for  months,  resisting  all  remedies,  and  often  accompanied  by  colicky 
pains  and  symptoms  of  peritonitis.  The  stools  often  have  blood 
mixed  wdth  the  fluid,  very  glassy,  jelly-like  mucus,  more  or  less 
abundantly  as  in  the  cases  of  Moesasca,  Moosbrugger,^  Kahane,^ 
Girard,^  Poledne,'  and  Rippe.^  Nausea  and  vomiting  are  rarer 
symptoms. 

Diagnosis  as  a  rule  can  only  be  made  by  microscopical  examina- 
tion of  the  stools ;  together  with  the  eggs,  regular  and  beautifully 
formed  Charcot-Leyden  crystals  occur. 

The  prognosis  is  unfavourable  in  severe  infections,  in  slighter 
cases,  where  only  a  few  worms  are  present,  the  danger  of  important 
symptoms  is  less.  Treatment  consists  in  administration  per  os  of 
vermicides  and  in  local  treatment  of  the  large  gut.  A  remedy  which 
was  once  much  used  was  calomel,  which  is  much  lauded  by  Gibson 
and  given  as  follows  :  calomel  o'c6  grm.,  rheum.  o'3  grm.,  tinct.  ferri 
sesquichlor.  1-2  c.c,  aq.  dest.90'o  grm.,  six  dessert-spoonfuls  three  times 
daily.  Rippe  appears  to  have  got  no  result  from  the  use  of  this  pre- 
scription.    Thymol,  especially  in  conjunction  with  local  treatment  of 


'  Moosbrugger,  Med.  Corresp.-BL  f.  Wilrttemburg,  1890. 

^  Kahane,  Korrespondenzhl.  f.  Schweiz.  AeztCy  1907,  viii. 

^  Schulze,  Deutsch.  med.  Wochenschr.^  1905. 

^  Vix,  Zeitschr.  f.  Psychial.,  xvii.  ^  Girard,  Annal.  d.  VInst.  Fasieur^   1901. 

^  Blanchard,  Acad,  de  Mid.^  July  3,  1906. 

"^  Poledne,  Wien.  med.  Wochenschr.,  1906. 

^  Rippe,  St.  Petersb.  med.  Wocheiischr.,  1907. 


68o  THE   ANIMAL   PARASITES   OF   MAN 

the  large  intestine,  had  unquestionably  some  effect  in  certain  cases, 
such  as  those  of  Girard,  Poledne,  Hausmann,  Kahane  and  Schiller. 
The  local  treatment  of  the  large  bowel  is  most  effectual  when  high  in- 
jections of  water  and  benzine  are  given.  Becker^  obviously  used  too 
much  benzine  (i  dessert-spoonful  to  i  litre  of  water),  for  severe 
irritation  was  set  up,  whilst  Peiper^  used  only  a  few  drops  of  benzine, 
5  drops  to  I  litre  of  water  being  enough  (Schiller).  Instead  of 
benzine  enemata,  garlic,  i  per  cent,  thymol  solution,  and  physiological 
saline  injections  have  been  used,  but  the  benzine  enemata  seem  to  be 
far  and  away  the  most  effective.  In  Schiller's  case  2,000  worms  came 
away  on  the  first  day  as  the  result  of  such  a  combined  treatment 
(thymol  internally  and  benzine  enemata). 

Trichinella   spiralis. 

Trichinosis  is,  happily,  becoming  so  much  rarer  that  many 
doctors  get  no  opportunity,  either  in  their  student  days  or  in  private 
practice,  of  seeing  this  severe  disease ;  we  ourselves  remember  having 
observed  one  typical  case  of  a  peasant,  aged  17,  from  Metz  in  Med.- 
Rat  Merkel's  clinic  in  Nuremberg  in  the  year  1879.  In  the  description 
of  the  disease  we  follow  Merkel's'*   observations. 

The  eating  of  flesh  containing  Trichinie  is  often  followed,  if  not 
invariably  so,  by  gastric  disturbances  of  different  kinds,  especially  by 
vomiting  and  diarrhoea,  with  colic,  great  muscular  fatigue,  oedema 
of  the  eyelids,  muscular  swellings  with  hardness  and  extreme  painful- 
ness,  disturbance  of  ocular  movements,  of  deglutition  and  of  breathing, 
hoarseness,  aphonia,  intestinal  haemorrhage,  bleeding  of  the  nose, 
ecchymosisof  the  skin  and  mucosae,  prurigo,  herpes,  miliaria,  pustules, 
boils,  severe  sweating,  oedema  of  the  extremities,  and,  finally, 
desquamation  of  the  skin  ;  more  rarely  there  is  considerable 
decubitus,  bronchial  catarrh,  hypostatic  and  catarrhal  pneumonia, 
with  dry  and  purulent  pleurisy,  and  in  severe  cases  symptoms  of 
collapse  with  delirium  close  the  scene.  Slight  cases  last  from  three 
to  six  weeks,  severe  ones  for  several  months,  and  in  the  latter 
convalescence  is  very  slow.  It  is  remarkable  that  in  cases  of 
trichinosis  of  long  duration,  cancer  of  the  breast  was  observed  at 
the  same  time  (Klopsch,-  Langenbeck,^  Babes^).  Death  during 
epidemics  occurred  in  30  per  cent,  of  all  cases.     The  disease  begins 

'  Becker,  Deuisch.  med.  Wochenschr.,  1902. 
-  Peiper,  quoted  by  Seifert,  loc.  cit.y  p.  248. 
^  Merkel,  "  Handb.  d.  Therap.  v.  Pentzoldt-Stintzing,"  i. 

*  Klopsch,  quoted  by  Babes. 
^  Langenbeck,  ibid. 

*  Babes,  Centralbl.  f,  Bakt.,  1906,  xlii. 


SUPPLEMENT  68l 

generally  from  one  to  ten  days  after  eating  trichinous  flesh,  yet  there 
have  been  cases  noted  in^. which  the  disease  began  several  v^reeks  after. 

Diagnosis  in  the  presence  of  several  cases,  or  in  epidemics,  is  not 
difficult,  but  in  isolated  cases,  on  the  other  hand,  it  is  not  easy. 
If  there  is  a  suspicion  of  trichinosis,  from  the  muscular  fatigue  and 
the  oedema  of  the  eyelids,  the  diagnosis  can  be  made  by  excision  of 
a  piece  of  muscle  and  by  finding  the  Trichinae  in  the  tissue,  taken 
with  the  results  of  the  examination  of  the  previously  eaten  sausage  or 
meat.  In  contradistinction  to  this  circumstantial  process,  there  is 
the  examination  of  the  blood,  which,  according  to  Schleip' 
(Homburg  trichinosis  epidemic,  August  19  to  26,  1903,  130  cases),  is 
the  most  valuable  method  of  diagnosing  trichinosis  when  the  Trichinae 
have  not  yet  penetrated  the  muscles,  for  a  blood  examination  shows 
a  large  increase  in  the  numbers  of  the  eosinophile  cells;  Staubli 
detected  his  seven  cases  in  this  way,  four  of  the  severe  on  s  showing 
a  marked  hyperleucocytosis,  and  a  combination  of  Kernig's  sign  with 
absence  of  the  patellar  reflex.  On  account  of  the  rarity  of  these  two 
signs  in  combination  in  other  infective  diseases,  they  have  a  certain 
diagnostic  value.  Staubli^  also  observed  in  trichinosis  the  constant 
appearance  of  a  remarkably  strong  positive  diazo-reaction  of  the 
urine. 

Prophylaxis  in  trichinosis  is  fully  considered  under  Tric/iinella 
spiralis  (p.  429). 

Treatment  consists  in  those  cases  where  it  is  known  that  trichinous 
flesh  has  been  swallowed  in  the  first  place  of  washing  out  the 
stomach,  but  still  more  in  a  thorough  evacuation  of  the  bowels,  for 
which  calomel  (0*5  grm.),  ol.  ricini  (a  dessert-spoonful  till  the  action 
becomes  marked),  infusion  of  senna  with  sulphate  of  magnesia  and 
large  enemata  are  employed,  and  should  be  repeated  at  intervals 
during  the  first  few  weeks.  Alcohol  (cognac  up  to  250  c.c.  a 
day)  is  recommended  by  some,  also  glycerine  (150  grm.  at  a  dose) 
and  large  doses  of  dilute  hydrochloric  acid.  Beside  these,  a  large 
number  of  other  remedies  are  recommended,  of  which,  perhaps, 
benzine  and  thymol,  especially  in  the  form  of  enemata,  are  worthy 
of  notice. 

When  the  disease  is  fully  developed  the  treatment  should  be 
symptomatic  ;  a  protracted  practically  continuous  luke-warm  bath  is 
especially  useful. 

Eustrongylus  gigas. 

Eustrongylus  gigas  is  most  frequently  found  in  the  pelvis  of  the 
kidney.     Infection   m  the  majority  of   cases  leads  to  pyelitis.     The 

^  Schleip,  Deutsch.  Arch,  f,  klin.  Med.^  Ixxx. 
^  Staubli,  ibid.,  Ixxxv. 


682  THE   ANIMAL   PARASITES   OF   MAX 

inflammation  extends  to  the  capsule  from  the  pelvis,  resulting  in  a 
purulent  nephritis.  In  infections  of  longer  duration,  the  affected 
kidneys  become  changed  into  so-called  kidney  sacs,  while  the  kidney 
itself  continuously  shrinks.  Owing  to  the  worm  fixing  its  posterior  end 
in  the  ureter,  and  owing  to  an  inflammatory  swelling  of  the  mucosa 
of  the  ureter,  the  passage  of  urine  becomes  very  difficult. 

The  symptoms  resemble  those  caused  by  a  foreign  body,  e.g., 
kidney  pain,  suppression  of  urine,  dysuria^  discharge  of  blood  and 
pus  with  the  urine.  But  these  symptoms  are  not  sufficient  for 
a  diagnosis  ;  this  can  only  be  established  by  finding  eggs  or  the 
parasite  itself  in  the  urine. 

Moscato^  records  a  case  with  chyluria,  pain  in  the  region  of  the 
right  kidney,  and  hysterical  symptoms.  During  an  hysterical  attack 
a  specimen  of  Eiistrongylus  gigas  was  discharged  in  the  urine,  and  the 
chyluria  and  nervous  aftections  disappeared.  In  a  case  described 
by  Stuertz-  of  an  Australian  with  chyluria  due  to  Eiistrongylus  gigas 
the  chyluria  had  existed  for  seven  years.  In  the  urine  the  eggs  of 
Eustrongy Ills  gigas  were  found.  The  cystoscopic  examination  showed 
that  turbid  urine  was  discharging  from  the  left  ureter.  Nephrectomy 
was  considered. 

Ancylostoma  duodenale  (Ancylostomiasis). 

Whilst  up  to  quite  modern  times  it  has  been  generally  maintained 
that  the  great  majority  of  worm  diseases  cause  more  or  less  marked 
symptoms,  the  exact  investigations  of  the  last  few  years  have  made 
it  plain  that  the  great  majority  of  people  w^th  worms  are  not  only 
perfectly  healthy,  but  the  most  careful  clinical  observations  show  no 
single  sign  of  any  ill-effect  of  the  intestinal  parasites  on  the  health 
of  the  host  (Lobker  and  Bruns^).  If  infection  has  led  to  the 
development  of  only  a  few  ancylostomes,  then  injury  to  the  general 
health  is,  as  a  rule,  scarcely  noticeable.  In  order  to  produce  severe 
illness  the  presence  of  several  hundred  worms  in  the  intestine  is 
necessary,  and  in  general  the  intensity  of  illness  varies  in  exact 
proportion  to  the  number  of  worms.  Then  the  duration  of  the  infec- 
tion comes  into  play  :  the  longer  the  human  organism  is  submitted 
to  the  injurious  effect  of  the  parasite,  the  clearer  is  the  effect  on  the 
host.  Besides,  the  resistance  of  the  individual  has  to  be  considered. 
Whilst  a  more  robust  person  can  harbour  without  ill-effect  for  a 
longer  time  a  larger  niunber  of  ancylostomes,  the  symptoms  of  the 
disease  become  more  markedly  and  much  sooner  apparent  in  weakly 
persons  or  in  those  weakened  by  other  diseases. 

*  Moscato,  quoted  by  Predtetschensky,  Zeilschr.  f.  klin.  Med.,  xl. 

^  Stuertz,  Ges.  d.  Charile-Aerzte  in  Berlin,  June  26,  1902. 

^  Lobker  and  Bruns,  Atb.  axis.  devi.  kaiserl.    Gestmdheilsatfiie,  1 906,  xxiii. 


,  SUPPLEMENT  683 

The  first  symptom  is  disturbance  of  the  digestive  system  ;  more 
often  there  is  a  feehng  of  pain  in  the  epigastrium,  more  severe  upon 
pressure,  heartburn,  nausea,  vomiting  of  mucus  or  food  at  different 
times  of,  the  day  (occasionally  ancylostome  ova  have  been  found  in 
the  vomit).  Whether  the  eggs  which  reach  the  frontal  sinus  with  the 
vomit  can  develop  into  larvae  there  is  questionable,  but  the  records 
of  v.  Ziemssen^  and  Huppertz,^  to  the  effect  that  in  some  instances 
ancylostomes  have  been  discharged  from  the  frontal  smus,  are  of 
interest.  The  five  cases  recorded  by  the  latter  had  a  fatal  termination 
from  oedematous  swellings  of  the  face  with  severe  inflammation  of 
the  meninges.  The  tongue  is  furred,  and  extensive  catarrhal  stoma- 
titis and  ptyalism  are  recorded.  The  appetite  is  variable,  increasing 
or  diminishing,  there  is  loathing  of  nourishment  or  a  marked  longing 
for  acid  food  and  unripe  fruit,  whilst  ordinary  meals  are  rejected. 
At  first  there  is  often  constipation,  later  diarrhoea  with  abundant 
mucus,  and  often  blood  in  the  stools;  microscopically  eggs  and 
Charcot-Leyden  crystals  w^ere  found. 

In  the  further  course  of  the  disease  symptoms  due  to  increasing 
anaemia  predominate  ;  the  haemoglobin  of  the  blood  diminishes  from 
one-fourth  to  one-fifth  of  the  normal  (Baravalle^),  the  eosinophile  cells 
increase  considerably  (Boycott,^  Lohr^),  yet  in  regard  to  diagnosis 
eosinophilia  cannot  be  regarded  as  of  equal  value  to  a  microscopical 
examination  of  the  faeces  (Bruns,  Liefmann,  and  Meckel^).  The  dis- 
turbances of  the  circulatory  system  take  the  form  of  more  or  less 
severe  palpitation,  pain  in  the  region  of  the  heart,  quick  pulse,  oedema 
of  the  eyelids,  of  the  face,  of  the  lower  limbs,  and  even  of  the  whole 
body.  Disturbance  of  the  sexual  functions  (impotence,  irregular 
menstruation,  delayed  onset  of  puberty)  are  not  infrequently  observed. 

Infection  in  human  beings  takes  place  by  the  mouth,  if  uncleansed 
vegetables  are  eaten — in  Japan  especially,  where  human  faeces  are  used 
— and  articles  of  food  are  not  sufficiently  carefully  cleaned  (Inouye^, 
or  from  putting  food  into  the  mouth  wdth  dirty  hands.  Looss^  does 
not  think  that  drinking  water  is  dangerous  as  a  rule,  for  the  larvae 
sink  to  the  bottom  in  standing  water,  and  are  only  brought  to  the 
top  by  shaking.  Looss  has  done  most  valuable  service  by  discovering 
that  infection  can  arise  also  through  the  skin.  During  the  last  few 
years  so  many  authors   have  confirmed  this  at  first   doubted  source 


1  V,  Ziemssen,  quoted  by  Haenisch,   "  Dis^.  Strasburg,"  1901. 

-  Huppertz,  quoted  by  Haenisch,  "Diss.  Strasburg,"  1901. 

■•*  Baravalle,  Progresso  medico^  I903' 

^  Boycott,  Journ.  0/ Hygiene,  1904.  '"  Lohr,  Zeitschr.  f.  Heilk.^  xxvi. 

6  Bruns,  Liefmann  and  Meckel,  Milnch.  med.  Wochenschr.,  1905. 

■^  Inouye,  Arch.  f.   Verdamcngs  Krankh.,  1905,  xi. 

^  Looss,  "  Handb.  f.  Tropenkrankh.,"  v.  Mense,  i,  p.  129. 

43 


684  THE   ANIMAL   PARASITES    OF   MAN 

of  infection,  that  one  must  accept  this  souice  of  infection  now,  even 
though  it  is  undecided  which  mode  of  infection  is  the  more  prevalent, 
by  the  mouth  or  through  the  skin.  Some  authors  have  described 
the  changes  induced  in  the  skin  by  the  penetration  of  the  larvae  ;  for 
instance,  Looss  and  Schaudinn,^  itching  papules  in  their  own  skin, 
and  Dieminger"^  a  skin  affection  in  the  Graf  Schwerin  mine  which 
was  called  the  "  Schweriner  itch,"  and  a  skin  affection  not  unlike 
scabies  in  the  tea  plantations  of  Assam  and  South  America  ;  pani- 
ghao  (water  itch)  (Dubreuilh^)  ;  the  penetration  of  the  larvae  through 
the  skin  also  explains  the  frequent  appearance  of  boils  and  itching 
purulent  eczema  in  miners  in  infected  pits  (Goldmann^). 

The  absolute  diagnosis  of  ancylostomiasis  depends  on  the  detection 
of  the  ancylostome  eggs  in  the  faeces,  and  presents  no  difticulties. 

Prophylaxis  is  of  the  greatest  importance,  especially  to  miners. 
The  spread  of  ancylostomiasis  seems  to  depend  only  on  faices 
deposited  in  damp  places,  so  that  on  the  one  hand  the  deposition 
of  faeces  must  be  prevented,  and  on  the  other  the  faeces  must  be 
rendered  as  far  as  possible  harmless  ;  in  addition,  there  is  the 
individual  prophylaxis. 

General  prophylaxis  requires  : — 

(i)  Examination  immediately  for  ancylostomes  of  miners  seeking 
work  and  of  those  newly  taken  on  five  to  six  weeks  after. 

(2)  Indentured  workers  who  are  infected  with  worms  are  not 
allowed  to  work  underground  until  a  medical  certificate  in  writing 
is  brought  to  the  effect  that  they  are  no  more  infected  with  eggs  (the 
same  procedure  applies  to  workmen  in  brick  kilns)  (Goldmann^). 

(3)  Indentured  workers  infected  with  worms  must  submit  them- 
selves to  the  prescribed  treatment,  and  after  its  completion  further 
submit  their  stools  to  three  examinations  at  intervals  of  about  four 
weeks. 

(4)  Special  supervision  of  miners  and  brick-makers  coming  from 
the  Italian  frontier. 

(5)  Workmen  must  be  given  instructions,  both  by  word  of  mouth 
and  in  writing  in  their  mother  tongue,  as  to  the  infectivity  and  danger 
of  ancylostomiasis  both  to  themselves  and  others. 

(6)  Orders  are  to  be  given  as  to  washing,  baths,  and  changing 
of  clothes  at  the  end  of  the  work. 

(7)  During  the  hours  of  working  in  the  pits,  taking  of  food  is 
strictly  forbidden  without  thorough  and  entire  washmg. 


'  Schaudinn,  Deutsch.  vied.  Wochenschr.,  1904. 

2  Dieminger,  Klin.  Jahrb.,  1905,  xiv.  ^  Dubreuilh,  La  Presse  mid.,  1905,  xxx. 

•*  Goldmann,  Wien.  nud.  Presse^  1905,  ii. 

■'  Ibid.^  "  Die  Hygiene  des  Bergmannts."  Halle  :   W.  Knapp,  1903. 


SUPPLEMENT  685 

(8)  All  privies  must  be  so  arranged  that  the  vessels  used  for  the 
reception  of  the  excreta  must  not  leak,  must  be  protected  by  a  cover, 
and  easily  transportable.  The  emptying  of  these  vessels  must  be 
carried  out  in  specially  constructed  impenetrable  pits. 

(9)  Defaecation  in  any  other  place  than  a  privy  is  forbidden  (alike 
for  miners  and  brick-makers). 

(10)  The  manure  of  horses  used  in  the  mines  is  to  be  regularly 
removed  ;  possibly  infection  takes  place  in  this  way  also.  [This  is 
impossible.— J.  W^  VV.  S.] 

How  far  it  is  possible  to  disinfect  a  mine  ah-eady  severely  infected 
is  a  matter  of  question  ;  Tenholt,^  Goldmann,'-^  and  Dieminger* 
recommend  washing  out  with  freshly  prepared  lime  water  witii  the 
addition  of  caustic  soda  ;  Calmette*  and  Manouriez^  spraying  with 
salt  water.  Theoretically  spraying  with  hot  water  or  steam  should  be 
done  every  now  and  again  for  the  destruction  of  the  larvae  (Looss*^), 
Personal  prophylaxis  is  partially  included  in  the  general  prophylaxis 
in  so  far  as  it  is  a  case  of  oral  infection,  but  something  more  can  be 
done  for  the  individual  to  avert  the  danger  of  cutaneous  infection. 
According  to  Manson'  it  is  advisable  in  the  tropics  to  cover  the 
naked  hands  and  feet  with  green  Barbados  tar,  and  the  tarred  parts 
thickly  with  flour;  Fabre^  recommends  that  miners  who  might  come 
in  contact  with  infected  water  should  anoint  the  unprotected  parts 
(hands  and  feet),  as  then  the  larvae  cannot  penetrate  the  skin  ;  this  last 
procedure  can  easily  be  carried  out  on  account  of  its  simplicity  and 
cheapness. 

Among  the  usual  remedies  for  the  expulsion  of  ancylostomes 
thymol  certainly  comes  first,  introduced  by  Bozzolo^  and  since 
used  by  many  other  authors,  partly  with  good  and  partly  with  less 
good  results.  The  day  before  the  beginning  of  treatment  one  should 
endeavour  to  procure  a  thorough  evacuation  of  the  bowels  by  means 
of  calomel  (Lutz,^^'  Griinberger,"  Smith^^)  or  cascara  sagrada  (Mann^^), 
only  fluid  food  should  be  taken  the  evening  before,  and  on  the  day 
of  treatment  thymol  is  given  in  a  quantity  of  6,  8,  10  or  15  grm.,  in 
single  doses  of  2  grm.  with  one  or  two  hours'  interval,  and  some  hours 
after  an  aperient.     As  a  rule,  one  day  of  this  treatment  is  not  enough 

'    Tenholt,  Miinch.  med.  Wochenschr.^  1905. 
-  Goldmann,  Wien.  med.  IVochenschr.,  1905,  x. 

=*  Dieminger,  loc.  cit.  ^  Calniette,  Acad,  de  Med.^  July  25,  1905. 

^  Manouriez,  Bidl.  de.  V Acad,  de  Med.,  1905. 
^  Looss,  Zeitschr.  f.  kliu.  Med.,  1905,  Iviii. 

'  Manson,  Brit.  Med.  Jown.,  November  5,  1900.  ^  Fabre,  Progrcs  meJ.,  1905. 

^  Bozzolo,  Giorn.  del  R.  Acad.  d.  Med.  dl  Torino,  1881.        '"  Lutz,  Centralbl.  f.  Bakt. 
■''  Giunberger,  Wien.  mei.  Wochenschr.^  1902,  lii. 
•■^  Smith,  Amer.  Joiirn.  Med.  Sci.,  1903. 
'^  Mann,  Deutsch.  Arch.f.  k'.in.  Med.,  Ixxiv. 


686  THE   ANIMAL   PARASITES   OF   MAN 

(Prowe^),  but  one  is  compelled  to  repeat  it  on  two  consecutive  days, 
or  even  oftener,  with  subsequent  intervals  of  many  days.  Thymol  is 
either  given  in  wafers,  gelatine  capsules  or  mixed  with  sugar.  Caution 
should  be  used  in  giving  brandy  at  the  same  time  or  bodies  which 
dissolve  thymol  (oil,  fat)  and  thereby  considerably  favour  its  absorp- 
tion. It  has  been  shown  in  many  cases  from  toxic  phenomena 
that  thymol  is  by  no  means  an  indifferent  drug;  violent  burning  in 
the  stomach  and  alimentary  canal,  lowering  of  the  temperature, 
shortness  of  breath  and  feeble  pulse,  giddiness,  delirium  and  fainting 
have  all  been  observed.  Sandwith^  and  Thornhill.^  as  well  as 
Leichtenstern,^  even  record  cases  of  death  after  the  use  of  thymol ; 
4  grm.  thymol  caused  severe  symptoms  of  poisoning  in  Griinberger's^ 
case.  The  black  colour  of  the  urine  (thymoluria)  which  so  often  sets 
in  after  the  first  dose  is  quite  harmless,  and  is  no  contra-indication 
to  the  continuance  of  the  cure.  Now^  and  again  there  are  traces  of 
albumin  in  the  urine,  but  it  is  very  seldom  there  is  any  severe  acute 
inflammation  of  the  kidneys.  Thymol  is  contra-indicated  in  advanced 
old  age  and  in  debility,  also  in  cases  with  a  tendency  to  vomiting,  in 
gastritis,  dysentery,  heart  or  kidney  affections. 

The  combination  recommended  by  Goldmann^  under  the  name 
of  taeniol,  already  mentioned  under  the  treatment  of  tapeworms,  and 
which  consists  of  thymol,  sebirol  and  salicylate,  appears  also  to  render 
good  service  in  the  treatment  of  ancylostomiasis  (Goldmann^  and 
Liermberger®). 

A  carbonate  of  thymol,  thymotal,  from  which  thymol  separates  off 
in  the  intestine,  is  given  three  to  four  times  a  day,  in  doses  of 
3  grm.  per  diem  (children  up  to  i*o  grm.)  on  four  consecutive  days, 
and  at  the  end  of  the  treatment  a  purge  (Pool,^  Bauer^^) ;  Leonardi^^ 
speaks  well  of  thymol  essence  (4*0  c.c.  per  diem)  in  an  emulsion  with 
plenty  of  water. 

The  next  drug  for  the  expulsion  of  ancylostomes  is  extrnctum 
iilicis  maris,  which  is  to  be  employed  as  in  tapeworm  treatment,  but 
has  not  always  had  the  desired  result,  whilst  in  such  cases  as  resist 
the  fern  extract,  thymol  attains  the  desired  effect  (Mann^^),  whilst  the 
reverse  is  frequently  observed  (Grunberger^^).  Nagel^'*  prescribes 
extr.  fil.  8  to  10  grm.,  chloroform  10  to  15  drops,  syr.  sennae  16  grm. ; 
before  taking,  the  glass  must  be  placed  in  hot  water,  otherwise  the 

'  Prowe,  Virch.  Arch.,  clviii.  '^  Sandwith,  quoied  by  Luubs. 

3  Thornhill,  ibid.  *  Leichtenstern,  Deutsch.  ined.  Wochenschr,^  1887. 

•'  Grvlnberger,  loc.  cit.  ^  Goldmann,  Ges,f.  inner e  Med.  in  Wien^  March  8,  1 906. 

'^  Goldmann,  Wien.  fued.  IVochenschr.,  1905,  x. 

"  Liermberger,  Berl.  klin.  Wochenschr.,  1 905. 

^  Pool,  Med.  Woche,  1901.  '«  Bauer,  Wien.  klin.  Wochejisckr.,  1904. 

"  Leonardi,  Gaz.  d.  Osp.,  1904.  '-  Mann,  loc.  cit. 

18  Griinberger,  loc.  cit.  '^  Nagel,  Deutsch.  vied.  Wochenschr.,  1903. 


SUPPLEMENT      687 

contents  will  not  pour  freely.  Zinn^  prefers  extract,  filicis  maris 
(freshly  prepared)  to  all  other  drugs.  Warburg'^  considers  the  treat- 
ment with  extr.  fil.  to  be  all  the  more  certain  the  more  thoroughly  the 
preliminary  treatment  is  carried  out.  Filmaron  07  grm.,  thymol 
5*0  grm.,  chloroform  i'5  grm,  ol.  ricini  20'o  grm.  gave  good  results 
after  being  given  two  to  three  times  (NageP).  Opinions  are  divided 
as  to  the  combination  of  thymol  and  extractum  hlicis  maris  (Hynek,"* 
Stockman,^  Boycott  and  Haldane,^  Adams'").  As  regards  other  remedies , 
eucalyptus  oil  is  well  spoken  of  by  Philips^  and  Hermann^  :  ol.  eucalypti 
2-0  grm.,  chloroform  3'o  grm.,  ol.  ricini  30'o  grm.,  to  be  taken  at  one 
time  or  in  three  separate  doses  in  the  morning  (on  the  previous  evening 
a  saline  purgative).  Neumann^^  recommends  podophyllin,  to  be  taken 
twice  on  three  consecutive  days  in  doses  of  0*035  Si''^"*-  Podophyllin 
appears  to  produce  quite  a  peculiar  condition  of  the  intestinal  mucosa 
which  is  very  prejudicial  to  the  Ancylostoma  adhering  to  it.  Bentley^^ 
regards  /3-naphthol  as  the  best  drug;  after  previous  examination  of  the 
bowels  he  gives  it  two  or  three  times  at  two-hourly  intervals,  in  doses 
up  to  ro  grm.  (Vide  also  the  Appendix,  p.  754,  for  other  drugs.) 
For  the  treatment  of  the  anaemia,  which  often  persists  very  obstinately, 
good  and  abundant  food,  iron  and  arsenic  preparations,  Levico  water 
(Goldman n,^"^  Liermberger^^)  are  suitable. 


Ascaris  lumbricoides  (Ascariasis). 

Ascaris  liunhricoldes  is  one  of  the  most  frequent  parasites  that 
occur  in  man,  both  in  adults  as  well  as  in  children  ;  as  a  rule,  indeed, 
it  most  frequently  infects  children  of  medium  age.  The  normal 
situation  is  the  small  intestine  ;  this,  however,  is  frequently  left,  and 
the  Ascarides  travel  into  the  stomach,  oesophagus,  pharynx,  bronchi^ 
the  nasal  cavities  and  still  other  regions.  It  is  a  peculiarity  of  the 
Ascarides  that  they  are  prone  to  glide  into  narrow  canals;  for  example, 
Clason^^  records  that  in  the  case  of  an  idiot  whose  custom  it  was  to 
swallow  glass  beads,  the  Ascarides  showed  a  predilection  for  sticking 
in  the  beads  and  were  passed  in  the  faeces.  The  disturbances  which 
Ascarides  occasion  in  the  intestine  itself  vary  ;  isolated  species  do  not 


'  Zinn,   "Theiap.  der  Gegenwart.,"  1903. 

2  Warburg,  Mi'uick.  nied.  Wochenschr.,  1904. 

^  Nagel,  loc.  cit.  ^  Hynek,  Sbornik  Klinikyy  v. 

^  Stockman,  Brit,  Med.  jfotcrn.,  1904. 

^  Boycott  and  Ilaldane,  Joiirn.  of  Hyg.^  ix. 

"^  Adams,  Arch,  of  Pediat.,  1901.  ^  Philips,  Lancet^  1906. 

^  Hermann,  La  vied,  moderjte,  1905.  ^°  Neumann,  Deutsch.  vied.  Wochenschr.^  1904, 

"  Bentley,  Indian  Med.  Gaz.,  1904. 

'-  Goldmann,  Deutsch.  Aerzte-Zeitg.,  1903.  '^  Liermberger,  loc.  cit. 

^*  Clason,  see  Seifert,  Deutsch.  med.  Zeitg.,  1885. 


688  THE   ANIMAL   PARASITES   OE   MAN 

give  rise  to  any  symptoms  at  all,  whereas  a  large  number  may 
eventually  give  rise  to  severe  local  symptoms,  or  those  of  a  toxic  or 
reflex  nature  which  have  been  discussed  in  the  General  Section. 

Among  the  local  symptoms  are  the  following  :  loss  of  appetite, 
excessive  appetite,  perverted  sense  of  taste,  foetid  breath,  sensitiveness 
to  pressure  over  the  abdomen,  colicky  pains  and  irregularity  of  the 
bowels.  The  appearance  and  state  of  health  suffer  ;  the  patients, 
children  in  especial  frequency,  become  remarkably  pale  ;  their  com- 
plexions undergo  rapid  change,  and  rings  of  grey  or  bluish-brown  are 
seen  about  the  eyes.  Children  may  become  so  reduced  by  this  rare 
condition,  enteritis  verminosa,  due  to  Ascarides  in  large  numbers,  that 
suspicion  of  the  existence  of  intestinal  tuberculosis  arises.  Emaciation 
to  a  skeleton,  excessive  meteorism,  and  evacuations  of  thin  gruel-like 
stools,  sometimes  blood-stained,  are  observed  in  these  cases.  Even  in 
the  case  of  adults,  chronic  uncontrollable  vomiting  with  severe 
inanition  due  to  the  Ascarides  has  been  observed.  When  the 
Ascarides  escape  spontaneously  per  aniim,  they  frequently  cause  an 
exceedingly  troublesome  irritation  in  the  anal  region  (pruritus  ani). 

The  most  disagreeable  symptoms  and  those  most  dangerous  to 
life  arise  from  the  migrations  of  Ascarides  when  they  invade  the  bile- 
ducts ;  no  inconsiderable  number  of  cases  of  this  kind  are  recorded 
in  the  literature  (summarized,  up  to  the  year  1901,  in  Sick's^  Disser- 
tation). Penetration  post  mortem  (or  shortly  before  death)  of  the 
worms  into  the  bile-ducts  cannot  be  considered  as  a  rarity;  the  laxity 
of  the  muscular  orifices  easily  allows  of  this  invasion  also  in  other 
directions  on  the  part  of  the  parasite  in  its  escape  from  the  body  of  its 
dead  host.  The  occurrence  of  the  worm  in  the  biliary  passages  in  the 
living  is  to  be  regarded  as  still  less  frequent,  but  nevertheless  often 
enough  according  to  the  records  in  literature.  Sick^  was  able  to 
collect  as  many  as  sixty-one  such  cases,  to  w'hich  he  added  two  further 
fresh  cases  from  the  Tubingen  clinic,  that  is,  from  the  material  pro- 
vided by  his  father.  In  the  year  1891  Borger'^  collected  fifty-nine 
•cases  relating  to  the  invasion  by  Ascaridce  of  the  bile-ducts  and 
passages,  and  Dauernheim's^  Dissertation  treats  of  this  question  as 
well.  A  further  case  of  Ascaris  in  the  ductus  choledochus  (chole- 
•dochotomy)  is  recorded  by  Neugebauer.'^  In  the  ca^^e  of  Schupper^ 
(woman,  aged  52),  all  the  biliary  passages  were  distended  and  filled 
with  fourteen  living  Ascaridiv  (perhaps  as  they  were  livijig  they  had 
not   led   to  a  septic  infection   of  the   biliary  passages) ;    in  the  case 


'  Sick,   "Diss.  Tubingen,"  1901.  -'  Sick,  ibid.,  1901. 

^  Borger,  "Diss.  Munchen,''  1891.  •*  Dauernheim,  "  Diss.  Giessen,'"  1900. 

■'  Neugebauer,  Arch.  f.  klin.   Chir.,  1903,  Ixx. 

^  Schupper,  Gaz.  d.  Os/'.,  1904,  xxxiii. 


SUPPLEMENT  6S9 

C(niiinunicated  by  Schiller,^  an  Ascaris  had  gained  access  to  the 
biHary  passages  after  an  operation  for  cholehlhiasis  (with  distension 
of  the  gall-bladder  and  formation  of  a  fistula)  ;  it  had  kept  itself  alive 
here  eighteen  days  and  was  extracted  from  the  fistulous  opening. 
Epstein"^  confirms  the  correctness  of  the  explanation  of  the  mark  of 
strangulation  in  an  Ascaris  in  Mertens'^  case  (in  a  woman,  aged  30, 
there  was  first  icterus,  later  ascites,  anasarca,  swelling  of  the  liver, 
then  the  discharge  of  two  dead  Ascaridcv,  one  of  which  exhibited  a 
constriction  somewhat  behind  its  centre  ;  after  that  there  was  rapid 
improvement  in  all  the  symptoms)  ;  in  his  case  there  was  icterus  in 
consequence  of  closure  of  the  ductus  choledochus  by  an  Ascaris. 
After  the  discharge  of  the  worm  the  symptoms  persisted  ;  one  of  the 
Ascaridcv  had  a  typical  strangulation  mark.  From  the  observation 
recorded  by  Vierordt*  it  follows  that,  without  doubt,  mature  females 
can  penetrate  into  the  liver  and  there  deposit  egi^s  ;  in  addition,  that 
such  egf^s  appear  exceptionally  to  undergo  segmentation.  A  unique 
feature  in  this  case  consisted  in  the  exclusive  discharge  of  immature 
worms  almost  regularly  throughout  an  interval  of  nine  weeks  ;  this 
cannot  be  explained  from  our  present  knowledge  of  the  biology  and 
pathology  of  the  Ascaridcv.  These  worms  clearly  make  their  w^ay  from 
the  intestine  outw^ards,  through  the  opening  into  the  duodenum  of 
the  common  bile-duct,  and  unquestionably  the  fully  developed 
Ascarides,  with  the  aid  of  their  conical  head  end,  are  enabled  gradually 
to  penetrate  the  wall  of  the  ductus  choledochus  (Quincke^),  and  gain 
access  to  the  gall-bladder,  the  hepatic  duct  and  its  branches. 

The  changes  in  the  biliary  passages  and  the  liver  are,  on  the  one 
hand,  the  mechanical  results  of  a  partial  or  total  obstruction  to  the 
flow  of  the  bile,  and,  on  the  other,  of  inflammatory  processes.  The 
blocking  of  the  common  bile-duct  and  of  the  trunk  of  the  hepatic 
duct  leads  to  the  well-known  symptoms  of  biliary  engorgement; 
protracted  continuance  of  this  condition  has,  as  its  sequela,  general 
distension  of  the  whole  biliary  system  and  degenerative  destruction 
of  the  liver-cells.  If  the  Ascaris  is  situated  at  some  other  part  of  the 
biliary  system,  its  presence  causes  a  partial  arrest  of  the  flow  of  bile, 
with  the  corresponding  sequelae.  Many  Ascarides  perish  in  the  ductus 
choledochus,  and  here  and  in  the  gall-bladder  they  may  supply  the 
nucleus  of  a  gall-stone  ;  deeper  in  the  liver  this  does  not  appear  to 
happen  ;  the  dead  Ascaridcv  here  undergo  a  kind  of  maceration,  dis- 
integrate, and  may  be  completely  absorbed;  in  many  cases  the  worms 

'  Schiller,  Beitr,  zur  klin.  Chir.,  1902,  xxxiv. 

-  Epstein,  Deutsch.  Arch.  f.  klin.  Med.,  1904,  Ixxxi. 

3  Martens,  Deutsch.  med.  Wochenschr . ,  1898,  xxiii. 

^   Vierordt,  Volkmann's  Samvil.  klin.   Vortr.,  No.  375. 

•'  Quincke,   *•  Nothnagel's  Spez.  Path.  u.  Therap.,"'  1899,  xviii. 


690  THli:    AXIiMAL    PAKASniiS    OF    MAX 

continue  to  live  for  a  very  long  tune  in  the  biliary  passages.  When 
the  worms  infect  the  biliary  passages  through  the  invasion  of  intes- 
tinal bacteria,  liver  abscesses  arise  (Daiiernheim/  Saltykow^). 
Leer^  goes  so  far  as  to  maintain  that  Ascaridiv  may  be  the  second 
most  frequent  cause  of  liver  abscesses.  That  Ascaris  in  the  pancreas 
may  simulate  liver  abscess  in  a  remarkable  fashion  is  shown  by 
Vierordt's^  observation,  which  is  quite  unique,  while  Ascaridcv  have 
been  found  to  occur  in  isolated  instances  in  the  excretory  ducts  of 
the  pancreas  and  in  its  branches,  where  they  have  remained  living  for 
a  long  time. 

It  is  no  rare  occurrence  for  Ascarichv,  in  consequence  of  their 
migration  into  the  stomach,  to  be  ejected  by  the  act  of  vomiting,  and 
in  such  way  to  gain  access  into  the  upper  air  passages,  or  to  find 
their  way  during  sleep  into  the  nose  or  accessory  sinuses  (Hosier  and 
Peiper'')  without  giving  rise  to  special  symptoms.  For  example, 
Troja*^  found  in  the  frontal  sinus  of  a  cadaver  a  large  coiled-up 
Ascaris  which  occupied  the  whole  cavity.  Wrisberg^  made  the  same 
observation  in  the  cadaver  of  a  boy.  Deschamps*^  and  Fortessin'^ 
mention  an  Ascaris  being  met  w^ith  in  the  antrum  of  Highmore. 
Observations  of  the  discharge  of  living  or  dead  Ascarides  from  the 
nose  are  frequently  recorded.  To  this  class  belongs  the  case 
mentioned  by  Albrecht,^*^  in  which  an  Ascaris  was  removed  from  the 
nose  of  a  girl,  aged  7  ;  also  the  case  recorded  by  Benievini,^^  from 
the  nose  of  one  of  whose  friends  a  worm  escaped ;  he  had  suffered 
from  the  most  violent  headaches,  fainting  fits,  dimness  of  vision  and 
vomiting;  after  the  escape  those  untoward  symptoms  disappeared. 
Similar  records  have  been  made  by  Forest,^^  Lanzoni,^^  Langelott,^^ 
Tulpe,!'^  Reisel,!^  Fehr,i^  Bruckmann,^«  Bahr,^^  Slabber,^^  Lange,^! 
and    Chiari.22     p^  rarer  case  is  that  recorded  by  Haffner,^^  that  of  a 

'  Dauernheim,  /oc.  cit.  '^  Saltykow,  Prag.  Zeitschr.  f,  Heilk.,  1900, 

^  Leer,  Brit.  Med.  Journ.^  1906.  ^  Vierordt,  loc.  cii. 

^  Mosler  and  Peiper,  "  Nothnagel's  Handb.,"  1894,  vi.  ^  Troja,  Napoli,  I77r. 

"^  Wrisberg,  see  Blumenbach,  Goitingen,  1907. 

^  Deschamps,  see  Blass,   "Diss.  Strasburg,"  1902. 

^  Fortessin,  j^.?  Bardeleben,  *'  Lehrb.  d.  Chirurgie,"  1875. 

'°  Albrecht,  Cominer.  Noricum.  T.  I.  Annul.,  1 739. 

"  Benievini,  "  Prol.  Anat.  d.  Sin.  front.,"  Gottingen,   1779. 

*2  Forest,  see  Tiedemann,  Mannheim,  1844. 

'8  Lanzoni,  idem. 

'^  Langelott,  idem. 

'^  Tulpe,  idem. 

'^  Reisel,  idem. 

^'  Fehr,  idem. 

'^  Bruckmann,  Covime?-.   Noric.^  1739- 

'9  Bahr,  idem.  20  Slabber,  idem. 

2'  Lange,  "  Blumenbach's  Med.  Bibl.,"  Goilingen,   1788. 

22  Chiari,  "  Krankh.  d.  Nase,"  1902. 

■^3  HafiFner,  Berl.  klin.  IVochenschr.,  1880. 


SUPPLEMENT  69I 

child,  aged  4,  in  whom  an  Ascaris  reached  the  nasal  cavity  through 
the  act  of  vomiting,  and  from  there  it  gained  access  through  the 
naso-lachrymal  duct  and  the  inferior  lachrymal  sac  into  the  lower 
punctum  lachrymale,  from  which  half  of  it  protruded. 

Among  the  rarer  causes  of  the  occurrence  of  strange  bodies  in  the 
pharynx  and  naso-pharyngeal  cavity,  Jurasz^  mentions  in  the  first 
place  vomiting,  which  may  afford  opportunity  for  the  more  solid 
bodies  of  the  stomach  contents,  and  even  parasites  of  the  digestive 
tract,  especially  Ascaridcv,  to  become  firmly  lodged  in  the  pharyngeal 
or  naso-pharyngeal  cavity.  Ascaridcc  may  obtain  access  from  the 
naso-pharyngeal  cavity  to  the  middle  ear  by  way  of  the  Eustachian 
tube,  as  has  been  observed  by  Reynolds'^  and  Wagenhauser^ ;  in  the 
case  recorded  by  TurnbuU  (girl,  aged  8,  with  pains  in  her  ear)  the 
Ascaris  apparently  reached  the  external  auditory  meatus  by  the  same 
route. 

The  irritation  of  the  larynx  and  air  passages  by  Ascaridcv  is  far 
more  dangerous  than  their  penetration  into  the  nose  and  naso- 
pharyngeal cavity,  because  not  only  are  attacks  of  suffocation,  but 
sudden  suffocation  thereby  induced.  Oesterlein^  records  a  fatal 
attack  of  choking  from  Ascaridcc  in  the  trachea.  In  a  case  recorded 
by  Smyly^  of  a  boy,  aged  3^,  tracheotomy  for  extreme  asphyxia  was 
performed  without  relief.  At  i\\Q post-mortem  iho.  cause  of  the  asphyxia 
was  found  to  be  an  Ascaris  in  the  trachea.  Furst"  collected  twenty- 
five  observations  of  invasion  of  the  larynx  and  trachea  by  Ascaris. 
Mosler"^  reports  the  case  of  a  patient  with  aphonia  and  dyspnoea 
from  whose  larynx  an  Ascaris  was  removed.  Donati'-'  reports  a  case 
of  four  Ascarides  in  the  larynx,  and  Cerchez^*^  of  asphyxia  from 
Ascaridcs  in  the  larynx  or  trachea.  Wagner"  records  the  case  of 
a  boy,  aged  8,  in  whom  a  coil  of  worms  was  ejected  from  the  stomach 
by  vomiting  ;  the  mass  blocked  the  entrance  to  the  larynx  and  led  to 
death  from  suffocation.  A  case  similar  to  that  recorded  by  Smyly  is 
communicated  by  Rabot^^;  it  was  that  of  a  child  who  underwent 
tracheotomy  for  diphtheria,  and  who  was  not  relieved  by  the  opera- 
tion ;  when,  however,  an  Ascaris  appeared  in  the  cannula  and  the 
parasite  was  removed  the  child  breathed  well.  In  Negresco's^-^  case, 
that  of  a  boy,  aged  3,  an  Ascaris  gained  access  to  the  larynx  and  from 
there  into  the  trachea,  and  a  fatal  issue  from  asphyxia  resulted. 

'  Jurasz,  Heymann's  "  Handb.  d.  Laryngol.  u.  Rhinol.,"  iii. 

-  Reynolds,  Lancet,  1880.  *  Wagenhauser,  Arch.  f.  Ohrenheilk.,  1889,  xxvii. 

^  Turnbull,   Vhchoxu-Hirsch  Jahresberickt,  18S0. 

"  Qe%\.tx\e\n  Deutsch.  Klin.  185 1.  "  Smy\y,  Dubl.  Journ.,  1867. 

'^  Furst,  Wieii.  vied.   PVochenschr.,  1879.  ^  Mosler,  quoted  l>y  Liesen. 

^  Donati,  Ann.  Univ.  de  Mid.  et  Chir.,  Milano,  1875. 

'"  Cerchez,  Clinica,  1891,  iv.  "   Wagner,  Dentsch.  med.   Wochenschr.,  1902. 

'-'  Kabot,  Soc.  de  Sci.  7n6d,de  Lyon,  September  9,  1904. 

'•'  Negresco,  Soc.  de  Med.  legale^  November  9,  1903. 


692  /    THE  ANIMAL  PARASITES  OF  MAN 

The  route  by  which  Ascaridcv  obtain  access  to  the  urinary  passages 
must  remain  undecided.  Schliiter*'  treated  a  woman,  aged  60,  with 
retention  of  urine.  Upon  catheterization  the  hinder  end  of  an 
Ascaris  hung  out  from  the  catheter  opening  ;  the  anterior  end  was 
fixed  in  tlie  tube  and  the  hunen  was  obstructed.  Perliaps  in  the 
female  sex  Ascaridcv  travel  from  the  gut  into  the  vulva  and  from  there 
into  the  bladder,  as  they  have  already  been  observed  in  the  vagina, 
where  they  cause  troublesome  symptoms  (pruritus  pudendi). 

The  diagnosis  of  ascariasis  is  not  in  general  difficult ;  now  and 
then  the  worms  are  discharged  spontaneously;  if  not,  the  ova,  which 
cannot  be  mistaken,  can  easily  be  detected  in  the  faeces  upon  micro- 
scopical examination.  Epstein's'^  method — namely,  on  every  occasion 
to  obtain  fresh  material  for  examination — is  much  to  be  recommended. 
This  consists  in  introducing  a  Nelaton's  catheter  into  the  rectum  with 
a  rotatory  motion  and  then  drawing  it  out.  A  small  portion  of  faeces 
forced  into  the  catheter  opening  is  more  than  sufficient  to  demon- 
strate the  presence  of  ova  of  the  parasites  upon  microscopical 
examination   of  a  preparation. 

In  spite  of  all  pressure  on  the  part  of  relatives,  treatment  directly 
against  Ascaridiv  should  not  be  carried  out  until  the  diagnosis  is 
certain. 

As  regards  prophylaxis,  much  can  be  done  l')y  not  throwing  the 
worms,  when  expelled,  on  to  the  dung-hill  or  into  the  privy,  but 
straightway  into  the  fire.  Metschnikoff"^  has  issued  a  warning 
.against  the  consumption  of  unboiled  or  badly  washed  vegetables, 
salad,  strawberries,  etc.,  and  also  against  drinking  polluted  water. 

For  the  expulsion  of  the  worms  flores  cinae  were  formerly  con- 
sidered the  most  useful  means ;  now,  however,  santonic  lactone — 
santonin — which  is  prepared  from  them,  is  almost  universally 
preferred.  By  many,  especially  in  practising  among  children,  flores 
cinae  are  still  recommended  in  the  form  of  Stork's  worm  electuary 
(consisting  of  flores  cinae,  rad.  jalapae,  valerian  and  oxymel  simplex). 
Guermonprez^  recommends  them  because  he  thinks  that  santonin 
only  excites  the  worms  and  consequently  causes  unpleasant  sym- 
ptoms. Besides,  in  the  form  of  the  above-mentioned  electuary,  flores 
cinae  can  also  be  given  several  times  daily  with  raspberry  jelly  up  to 
0-5  grm.  to  2  grm.  (children  and  adults). 

Santonin  is  prescribed  either  in  single  doses  from  0*03  to  0-05  to 
o-i  grm.  with  sugar  in  the  form  of  powder,  or  else  in  oily  solution. 
When  given  in  the  latter  form  the  absorption  of  the  santonin  in  the 

'  Schliiter,  Munch,  med.   IVochenschr.,  1902. 

-  Epstein,  see  Seifert,   "  Lehrb.  d.  Kinderkrankh.,"  p.  273. 

=*  Metschnikoff,  Gaz.  hebd.  de  Med.  et  Chir.,  igoi. 

^  Guermonprez,  see  Seifert,  Detitsch.  med.  Zeitg.,  1885. 


SUPPLEMENT  693 

stomach  is  excluded  and  the  whole  quantity  introduced  is  thus 
enabled  to  reach  the  worms  in  the  intestinal  canal.  Kiichenmeister^ 
has  already  recommended  combination  of  santonin  with  ol.  ricini. 
Lewin,^  however,  states  that  ol.  morrhuae,  ol.  olivarum,  ol.  cocos 
and  ol.  cinae  can  also  be  taken.  In  prescribing  santonin  in  oily  solu- 
tion Henoch^  also  prefers  the  combination  with  ol.  ricini.  According 
to  Lewin's  direction  the  prescription  would  run  as  follows  :  — 

^     Santonin     ...         ...         ...         ...         ...     0*2  grm. 

01.  ricini.    ...  ...  ...  ...  ...     20*0  grm. 

01.  cinae  selh.         ...         ...         ...          ...     gtt.  iv. 

M.,d.s. 

S.,  one  tablespoonful  to  be  taken  iwo  to  three  times. 

If  the  patients  should  manifest  a  repugnance  to  castor  oil, 
Starke's  ricinus  paste  may  be  selected:  — 

^     Santonin      ...  ...  ...  ...  ...  o*2grm. 

01.  ricini     ...  ...  ...  ...  ...  20'0  grm. 

01.  cinoe  seih.  ...  ...  ..  ...  gtt.  iv. 

Sacch.  albi.  ...  ...  ...  ...  q.s. 

Pasta  mollis. 

S.,  to  be  used  for  two  days. 

If  necessary  the  first-mentioned  mixture  might  be  given  in  gelatine 
capsules.  Small  children  should  be  given  0*025  grm.  santonin  in 
warm  olive  oil  slightly  sweetened  with  sugar  (a  teaspoonful)  in  the 
morning ;  if  in  the  course  of  the  forenoon  specimens  of  Ascaris 
escape,  a  second  dose  should  follow  in  the  afternoon  about  two 
hours  after  the  meal.  Older  children  should  be  given  santonin  in 
combination  with  castor  oil  or  calomel  : — 

^     Santonini o*oi  to  o'02  to  0*03  grm. 

Calomelan  ...         ...         ...         ...         ...     0*025  grm. 

Sacch.  albi o*5gim. 

M.f.p.     D.  tal.  dos.  X. 

S.,  one  powder  about  six,  seven,  and  eight  o'clock  on  three  consecutive  days. 

As  santonin  causes  slight  toxic  symptoms  such  as  urticaria, 
vomiting,  retention  of  urine,  headache,  vertigo,  yellow  vision 
(xanthopsia),  it  is  in  every  case  advisable  to  follow  with  a  laxative  to 
expel  the  drug  from  the  body  as  speedily  as  possible.  The  urine 
is  coloured  yellow  from  one  to  two  days  and  assumes  a  scarlet 
red  colour  upon  the  addition  of  alkalis  ;  this,  however,  soon 
disappears,  while  it  persists  in  the  case  of  rhubarb  and  senna. 

In  the  place  of  santonin  iodoform  in  the  form  of  a  powder  mixed 


'  Kiichenmeister,  loc.  cil.  -  Lew  in,  see  Sti'ett,  Deut^ch.  vied.  Zeitg.^  1885. 

^  Henoch,  idem. 


694  THE    ANIMAL    PARASITES   OF   MAN 

with  bicarbonate  of  soda  is  given  by  Schidlowsky^  in  doses  up  to 
o*oi  to  o'o6  grm.  three  times  daily,  and  a  dose  of  castor  oil  on  the  day 
after  the  iodoform  is  given.  Thymol  in  addition  to  thymol  enemas 
may  be  tried,  in  doses  up  to  0*5  to  2-0  grm.  per  diem  (Calderone,^ 
Hausmann-^),  also  /3-naphthol  up  to  o'45  grm.  three  times  daily  (Du 
Bois^),  and — 

^j^;     Benzo-naphthi.l      2-ogrm. 

Semin  cinae...  ..  ...  ...  ...      logrni. 

Sacch.  albi o^grm. 

M.jf.p.     Divide  in  pait.  seq.  xxii. 

S.,  three  lo  five  powders  daily. 

(Ferran^),  tilmaron  oil  i'd  to  2-0  to  3*0  grm.  in  gelatine  capsules^ 
according  to  age  (Bodenstein^).  Bnining^,  ^  recommends  the  so-called 
American  worm-seed  oil,  derived  from  a  plant  native  to  the  United 
States,  Cheiiopodiuin  anfhdminticinn,  Gray.  It  is  given  in  emulsion 
(ol.  chenopodii  anthelm.  ico  grm.,  vitelli  ovi  unius,  ol.  amygd.,  gi. 
arab.  pulver.  aa  io*o  grm.,  aq.  destill.  200  grm.  ;  f.  emulsio)  up  to  0*25 
to  o*5  grm.  three  times  daily  at  one  to  two-hourly  intervals,  or  as  a  pure 
oil  from  8  to  15  drops  in  sugar  and  water  ;  to  be  followed  an  hour 
after  the  last  dose  by  oleum  ricini  or  pulvis  curellae.  If  no  action 
takes  place  by  the  afternoon,  a  laxative  should  again  be  given.  The 
treatment  frequently  must  be  repeated  the  next  day.  Thelen°  appears- 
to  have  had  good  results  from  this  drug. 

Corsican  moss  (mousse  de  Corse),  kamala,  Artemisia  absinthium, 
valerian,  semen  sabadillae,  have  all  been  supplanted  by  santonin  and 
at  most  are  used  as  adjuvants  for  the  latter. 

Oxyuris  vermicularis   (Oxyuriasis). 

Oxytiridce  do  not  remain  at  rest  in  the  gut,  but  leave  it,  generally 
at  night  time,  to  migrate  around  the  anus,  into  the  gluteal  folds,  and 
in  females  into  the  vulva  and  vagina  and  still  higher  up,  giving  rise 
in  these  different  sites  to  a  whole  series  of  irritative  symptoms.  In  the 
rectum,  also,  Oxyuridce  give  rise  to  such  symptoms,  which  are  mani- 
fested in  the  form  of  catarrhal  inflammation  ;  numerous  chronic 
intestinal  catarrhs  are  thus  explained.  The  frequent  coincidence  of 
haemorrhoidal  troubles  with  Oxyuridce  may  be  attributed  to  the  fact 
that  the  veins  of  the  rectum  participate  in  those  changes  which  have 


'  Schidlowsky,  see  Seifert.  -  Calderone,  idon. 

^  Hausmann,  St.  Peter sh.  vied.   Wochenschr.,  1900. 

^  Du  Bois,  see  Lenhartz  in  "  Penzoldt-Siintzing's  Handbuch,"  p.  619. 

^  Ferran,  idejn, 

^  Bodenstein,  Wien.  med.  Presse,  1906.  '  Bruning,  Med.  Klin.,  1906. 

**  IJemy  Deutsch.  med.  Woe  hens  chr.,  1907. 

^  Thelen,  *'  Diss.  Rostock,"  1907. 


SUPPLEMENT  695 

been  described  as  occurring  in  the  intestinal  mucosa.  Oxyuridcv 
may  also  give  rise  to  prolapse  of  the  anus,  either  by  the  tenesmus 
they  bring  about  having  such  a  prolapse  as  its  direct  sequel,  or  the 
proctitis  that  supervenes  constituting  a  further  etiological  factor  for 
its  occurrence  (Ungar').  Anal  fistulae  which  still  fiu-ther  increase 
the  trouble,  and  even  rectal  listulac;,  appear  to  be  capable  of  onset  in 
consequence  of  the  irritation  of  the  mucosa  brought  about  by  Oxy- 
jiridce  (Trendelenburg"^).  The  conditions  recorded  by  von  Wagener^ 
and  Ruffer^  appear  to  be  of  interest.  At  the  post-mortem  on  a  child, 
aged  5,  the  former  found  fifteen  to  twenty  quite  minute  nodules  on 
some  Peyer's  patches,  and  in  several  of  these  OxynrUhv  were  found 
upon  microscopical  examination  between  the  calcareous  concretions 
within  the  patches.  He  presumes  that  the  parasites  penetrated  the 
follicular  ulcers,  and  after  healing  of  the  latter  that  they  died  and 
became  calcified.  In  the  case  of  a  man  who  died  from  cirrhosis 
of  the  liver,  Ruffer  found  in  the  rectum,  at  a  distance  of  about  6  in. 
from  the  anal  orifice,  several  tumours  covered  by  the  intestinal 
mucosa,  the  smallest  of  which  was  the  size  of  a  pin's  head  and  the 
largest  that  of  a  w^alnut.  The  tumours  looked  like  calculi  overgrown 
by  connective  tissue  ;  under  the  microscope,  countless  Oxyuridcv  ova 
were  found  in  their  interior. 

The  symptoms  of  irritation  set  up  by  these  migrations  from  the 
intestine  are  troublesome  to  the  last  degree  ;  the  pruritus  thereby 
induced  is  often  unendurable ;  as  this  irritation  from  itching  comes 
on  with  especial  severity  during  the  night,  the  night's  rest  is  grievously 
interfered  with  ;  many  attacks  of  night  terrors  appear  to  be  occasioned 
by  these  worms.  But  the  general  condition  suffers  as  well  ;  the 
children  become  pallid  and  affected  with  nervous  excitability. 
Through  the  act  of  scratching  the  irritated  parts  the  ova  of  the 
parasites  may  be  conveyed  by  contaminated  fingers  directly  into  the 
oral  or  nasal  cavities,  certainly  also  into  the  oral  cavity  by  the 
contamination  of  food  (auto-infection).  In  the  case  of  boys  the 
sexual  organs  may  be  excited  sympathetically  through  irritation  of 
the  sacral  nerves  of  the  rectum  ;  girls  may  be  induced  to  practise 
onanism  in  consequence  of  the  entrance  of  the  worms  into  the  vulva. 

As  a  result  of  the  itching  irritation  which  the  scratching  gives  rise 
to,  and  of  the  irritation  due  to  the  parasites  migrating  to  the  area 
surrounding  the  anus,  congestion  and  inflammatory  symptoms  may 
arir^e  in  the  peri-anal  and  perineal  regions  (weeping  eczema,  Seifert),^ 


'   Ungar,  see  Seifert,  "  Lehrbucli  der  Kinderkrankh.,"  p.  246. 
-  Trendelenburg,  see  Seifert,  f'de/n. 

3  von  Wagener,  Deutsch.  Arch.  f.  Jdhi.  Med.,  Ixxxi. 

4  Rufifer,  Brit.  Med.Journ.,  1 90 1. 

*  Seifert,  '*  Lehrb.  d.  Kinderkrankh.,"  and  Lesser's  "  Encyklop.  d.  Haut-u.  Geschlechts- 
krankh.,"  p.  373. 


696  THE    ANIMAL   PARASI  lES   OF   MAN 

and  tliese  do  not  abate  till  after  the  removal  of  the  oxyuriasis.  Some 
authors  speak  of  an  oxyuriasis  cutanea  (Majoclii^),  in  the  more 
limited  sense  of  a  dermatitis  intertriginoides.  So  far  five  such  cases 
have  been  recorded,  one  each  by  Szerlecky/  Michelson,^  Majochi/ 
Barbagallo''  and  Vignolo-Lutati.^  Szerlecky's  case  was  that  of 
a  young  woman  with  intertrigo  over  the  thighs  (the  skin  was  covered 
as  if  with  leather)  ;  Michelson's  case  was  tiiat  of  a  boy,  aged  13, 
with  intertrigo  on  the  skin  of  the  genito-crural  fold,  of  the  scrotum 
and  of  the  thigh  ;  Majochi's  was  that  of  a  man,  aged  38,  with  the 
same  localization  ;  Barbagallo's  case  was  that  of  a  boy,  aged  14, 
in  vviiom  the  dermatitis  extended  to  the  hypogastrium  (rhagades  on 
the  scrotum)  ;  and  Vignolo-Lutati's  case  was  that  of  a  man,  aged  24, 
with  intertrigo  of  the  peri-anal  and  perineal  I'egion,  of  the  scrotum 
and  the  inner  side  of  the  thigh. 

On  leaving  the  gut,  Oxyuridai  frequently  migrate  to  the  stomach, 
to  the  oesophagus,  to  the  mouth,  to  the  nasopharyngeal  cavity,  and 
into  the  nose  (Zarniko')  (the  localization  in  the  nose  has  been 
referred  to  as  associated  with  the  possibility  of  auto-infection — sec 
p.  695  as  to  the  development  of  embryos  from  the  ova  in  the 
moist  nasal  mucosa).  Still  the  occurrence  of  Oxyiiridcv  in  the  nose 
is  among  the  greatest  of  rarities.  Chiari^  records  the  case  of  a  girl, 
aged  14,  who  suffered  from  pains  at  the  root  of  the  nose  and  in 
the  left  side  of  the  forehead;  female  specimens  of  Oxyurlsveniiicularis 
were  evacuated  from  her  nose  on  several  occasions.  A  similar  case 
is  recorded  by  Hartmann^ ;  it  was  that  of  a  girl,  aged  13,  with 
epileptiform  convulsions  and  psychic  disturbances  ;  numerous 
Oxyurides  frequently  escaped  from  her  nose.  With  their  departure 
the  symptoms  of  irritation  of  the  central  nervous  system  also  dis- 
appeared. Rheins^^  records  a  case,  that  of  a  woman,  in  which  a 
specimen  of  Oxyurls  vennicularis  was  discharged  from  the  right 
nostril  during  the  act  of  sneezing.  Proskauer^^  found  in  the  nose 
of  a  woman,  aged  30,  a  conglomerate  of  from  fifteen  to  twenty  very 
small  worms  which  proved  to  be  Oxyuris  embryos. 

The  diagnosis  of  oxyuriasis  is  not  difficult  to  make,  as  the  trouble- 
some sensations  in  the  anus  and  about  the  genitals  necessarily  suggest 
the   presence   of    Oxvnrldcv.     As  a  rule  the  small   white    worms  are 

'   Majochi,  Boll.  d.  Sci.  vied.  d.  Bologna,  1893. 
-  Szerlecky,  Jouni.  Ann.  Med.  praf.,  Paris,  1874. 
■'  Michelson,  Bej-l.  klin.   IVochenschr.,  1877,  xxxiii. 

*  Majochi,  loc.  cit.  ^  Barbagallo,  Gaz.  d.  Osp.,  November  16,  1900. 

•^  Vignolo-Lutali,  Atch.f.  De>-7ti.,  Ixxxvii,  pt.  I. 
'^  Zarniko,  "  Die  Krankh.  d.  Nase,  u.s.w,"  S.  Karger,  Berlin,  1905. 
"  Chiari,  "  Erfahr.  auf  d.  Gebiete  der  Hals-u.  Naseiikrankh.,"  Wien,  1887. 
®  Harlmann,  Natti'foischerversainmi.,  Koln,  1889. 
'"  Rheins,   "  Der  prakt.  Arzt.,"  1893.  "  Proskauer,  Zeituhr.f.  Ohienheilk.,  1891. 


SUPPLliMliNT  697 

seen  crawling  about  over  recently  evacuated  faeces,  or  the  ova  are 
found  upon  microscopical  examination  of  soiled  matter  adhering  to 
the  anus,  or  in  scrapings  removed  with  the  spatula  from  the  surface 
of  the  skin  (in  the  case  of  oxyuriasis  cutanea). 

Prophylaxis  has  to  be  directed  to  infection  with  Oxyurides 
generally,  on  the  one  hand,  and,  on  the  other,  to  the  possibility  of 
auto-infection.  With  reference  to  the  first-mentioned  point,  Metsch- 
nikoff's^  directions  should  be  borne  in  mind,  to  the  effect  that  badly 
washed  vegetables,  salad,  etc.,  ought  not  to  be  eaten  (vegetables  to  be 
rinsed  with  boiling  water),  and  also  that  the  members  of  the  family 
of  the  diseased  individual  should  be  examined  for  Oxyitridoe  and 
eventually  be  treated  (Heller^).  With  regard  to  the  second  point, 
one  has  to  observe  strict  cleanliness  in  general  (Barbagallo^  found 
ova  of  the  parasites  in  the  layer  of  dirt  under  the  finger-nails). 

Treatment  of  oxyuriasis  must  be  of  a  twofold  nature  ;  first, 
medicinal,  the  administration  per  os  of  vermicidal  drugs  in  combina- 
tion with  purgatives ;  and  secondly,  local  treatment  of  the  gut  by 
means  of  enemata,  suppositories  and  high  injections.  Following 
the  method  prescribed  by  Ungar,*  pulv.  glycyrrhizae  co.  is  first  given 
in  the  case  of  smaller  children,  castor  oil  or  calomel  m  that  of  those 
older,  in  order  to  evacuate  the  intestine,  and  four  times  daily  on  two 
days  following  one  another  a  dose  of  naphthalin,  not  du'ectly  after 
meal-time,  but  as  far  as  possible  in  the  interval  between  two  meals, 
and  at  the  same  time  the  ingestion  of  fatty  or  oily  nutriment  is  as 
far  as  possible  to  be  avoided.  After  eight  days  this  treatment  should 
be  repeated,  and  under  certain  circumstances  once  again  after 
a  further  interval  of  a  fortnight.  The  dose  varies  between  0-05  and 
01  grm.  (children  of  i  year  old),  0"i  toO'2grm.  (children  of  2  to  3  years 
old)  and  0*2  to  0*4  grm.  (children  of  4  to  10  years  old).  Dornbluth^ 
employs  the  same  medicament  in  a  form  only  slightly  modified  from 
Ungar's  method,  Barbagallo-  gives  internally  only  a  purgative  (decoct, 
sennae  cum  natr.  sulfur).  Thymol,  santonin,  kousso,  kamala  or  valerian 
may  be  tried  instead  of  naphthalin.  For  enemata  the  following  are 
employed  :  naphthalin  in  a  solution  of  i  in  50,  ol.  olivar.  or  thymol 
o*i  in  200  aq.  destill.,  diluted  solutions  of  lysol,  menthol  in  ^  per  cent, 
oily  solution,  salicylate  of  soda  in  watery  solution,  decoctum 
tannaceti  with  santonin,  with  the  addition  -of  some  drops  of  ol. 
terebinth.  (Barbagallo).  Decoctions  of  garlic,  infusion  of  valerian, 
sulphur  water  (sublimate  is  to  be  avoided),  aq.  calcariae,  ol.  olivanim 

'    Metschnikoff,  Med.  Klin.,  1907,  xlii,  p.  1284. 
-  Heller,  Deutsch.  Arch.  f.  klin.  Med.,  Ixxvii. 

*  Barbagallo,  loc.  cit. 

^  Ungar,  see  Seifert,  "  Lehrb.  d.  Kinderkrankh," 

•  Dornbluth,  Arzil.  Zenlral-Anzeiger,  1903. 
*5  Barbagallo,  loc.  cit. 


6g\<  THE   ANIMAL   PARASITIiS   OF   MAN 

cainphoratum  (Vignolo-Lutati).  Santonin  o'l  grm.  is  the  best  to 
employ  for  suppositories. 

For  high  injections,  large  quantities  of  plain  water  are  employed 
(2  to  4  litres),  or  soapy  water  (0*2  to  0*5  per  cent,  solution  of  sapo 
medicatus,  Heller,^  StilP),  J  per  cent,  salicylic  acid  solution  or  liq. 
alum.  acet.  (one  tablespoonful  to  a  litre  of  water,  Dornbliith^),  or 
gujanosol  (2  to  3  to  4  to  5  per  cent,  solution,  Rahn^).  The  employ- 
ment of  benzine  for  such  high  injections  is  not  advisable  according  to 
the  experience  of  Senger,-^  owing  to  the  symptoms  of  poisoning  after 
the  external  application  of  benzine,  at  least  not  in  the  case  of  young 
children. 

That  diseases  of  the  intestine  which  are  accompanied  by  frequent 
thin  fluid  evacuations  may  lead  to  recovery  from  oxyuriasis  has 
frequently  been  observed  by  us  in  the  case  of  young  children  who 
have  suffered  from  dysentery  (Seifert^).  Inunctions  of  cod-liver  oil 
appear  to  be  very  valuable  in  the  treatment  of  oxyuriasis  (Szerlecky, 
Vignolo-Lutati),  whilst  those  with  mercurial  ointment  may  easily 
increase  the  inflammatory  symptoms.  The  luxury  recommended 
by  Esser,^  that  patients  every  evening  before  going  to  sleep  should 
have  the  female  Oxyuridcv  picked  from  the  anal  fold  in  the  knee- 
elbow  position  is  one  which  is  certainly  only  in  the  power  of  a  few 
people  to  carry  into  execution. 

An  essay  has  been  published  by  Hippius  and  Lewinson  [Dsztlsch.  vied.  Wochensckr., 
1907,  xliii.)  in  which  the  relationship  of  Oxytcrida  to  appendicitis  is  considered  and  the  treat- 
ment of  oxyuriasis  is  discussed.  The  instructive  case  recorded  appears  to  show  that  germs 
through  Oxyicrida  gain  access  to  the  tissue  of  the  appendix,  and,  indeed,  are  carried  in  by 
them.  In  view  of  this  more  recent  communication  as  to  the  part  whicii  intestinal  parasites 
play  in  the  etiology  of  appendicitis,  it  seemed  to  me  [O.  S.]  to  be  worth  whi  e  to  interrogate  my 
surgical  colleagues  as  to  this  point.  About  2,000  appendicectomies  have  been  jointly  per- 
formed by  Drs.  Burkhardt,  Enderlen,  Pretzfelder,  Riedinger,  Rosenberger  and  Siber,  and  in 
not  one  of  these  cases  could  entozoa  be  found  to  be  a  possible  cause  of  the  appendicitis. 
Such  figures  without  doubt  speak  in  favour  of  the  fact  that  even  if  in  individual  cases  entozoa 
might  come  into  reckoning  as  a  possible  cause,  such  an  etiological  factor  must  be  classed 
among  the  greatest  of  rarities.  My  colleague,  Dr.  Ries,  who  practised  for  ten  years  in  Mexico, 
informed  me  that  there  practically  speaking  every  Indian  without  exception  harboured  para- 
sites of  the  most  varied  kind,  and  that  in  spite  of  the  very  extensive  professional  standing  he 
enjoyed  among  these  people  he  never  had  under  observation  among  them  a  single  case  of 
appendicitis.  As  far  as  the  observation  of  the  authors  in  question  as  to  the  treatment  of 
oxyuriasis  is  concerned,  it  must  be  energetically  directed  to  the  employment  of  local  measures 
for  the  intestine  ;  they  maintain  that  the  use  of  enemata  would  be  irrational,  and  that  it  is 
-astonishing  that  this  method  has  been  able  to  maintain  its  standing  down  to  the  present  day. 


'   Heller,  loc.  cii. 

-  Still,  Br  a.  Med.  Journ.,  1899.  ^  Dornbltith,  loc.  cit. 

*  Rahn,  Miinch.  med.   Wochenschr.,  1 905. 

*  Senger,  Berl.  klin.   Wochenschr.,  1907,  xxxviii. 
^  Seifert,  Deutsch.  med.  Zeilg.,  1885. 

"'■  Esser,  Schweiz.  Korrespondejizbl . ,  1 893- 


SUPPLEMENT  699 


HIRUDINEI    (Leeches). 

The  only  one  of  the  leeches  that  comes  under  consideration  from 
the  clinical  point  of  view  is  Limnatis  nilotica  [Hcemopsis  sanguisitga), 
which  obtains  access  to  the  mouth  with  drinking  water,  and  becomes 
lodged,  even  in  the  case  of  man,  in  the  pharynx,  larynx,  trachea, 
oesophagus  and  nose. 

Amongst  the  causes  of  severe  haemorrhage  from  the  pharynx 
Jurasz^  mentions  the  occurrence  of  leeches  in  that  region  :  in 
Northern  Europe  this  must  be  accounted  one  of  the  greatest  of  rarities, 
whilst  at  all  times  in  southern  countries,  such  as  South  Italy,  Spain, 
Greece,  Algiers,  Tunis  and  Egypt,  it  appears  to  have  been  more 
frequent.  Even  the  physicians  of  antiquity  had  much  to  say  about 
it.  Upon  the  occurrence  of  blood-stained  expectoration,  Hippocrates 
recommends  the  oral  cavity  to  be  examined  to  see  whether  a  leech 
is  not  present  in  it.  Galen  speaks  of  haematemesis  due  to  the  presence 
of  leeches  in  the  pharynx  and  stomach.  Similar  mention  is  found 
in  the  writings  of  Celsus,,  Asclepiades,  Scribonius  Largus,  Dioscorides, 
Aetius,  Oribasius,  Paulus  Aegineta  and  others.  In  recent  times, 
CortiaF  has  published  observations  relating  to  this  subject  which 
he  had  the  opportunity  of  making  in  Constantine.  Palazzolo^ 
also  in  Sicily  found  leeches  in  two  cases  in  the  pharynx,  in  one  case 
on  the  posterior  wall,  in  the  other  in  the  crypt  over  the  left  tonsil. 
According  to  Roset,*  leeches  adhere  by  preference  behind  the  uvula, 
simulating  haematemesis  and  haemoptysis,  and  the  persistent  haemor- 
rhages they  give  rise  to  may  lead  to  severe  anaemia.  Leeches  are  found 
in  still  greater  frequency  in  the  larynx  than  in  the  pharyngeal  cavity. 
Huber^  records  several  observations  of  this  kind  in  his  historical 
and  therapeutical  study.  In  the  case  of  a  man,  aged  64,  Ramon 
de  la  Sota  y  Lastra®  observed  a  leech  on  the  nodulus  epiglottidis  ; 
this  was  removed  with  the  forceps.  In  the  case  recorded  by 
Photiades,^  a  leech  had  remained  adherent  to  the  vocal  cord  for 
more  than  twenty-two  days.  Maissurianz^  records  two  such  cases : 
in  one  the  leech  had  remained  in  the  sinus  morgagni  for  three  weeks,, 
in  the  other  in  the  same  place  for  ten  days.     The  case  recorded  by 


*  Jurasz,  Heymann's  "  Handb.  d.  Laryng.  u.  Rhinol.,"  1899,  ii, 
-  Cortial,  Union  mid.^  1886. 

^  Palazzolo,  Bull.  del.  vial.  deW  orecchioy  etc,  1895. 

*  Roset,  Rev.  d.  Cienc.  niM.  de  Barcelona,  1 907,  ii. 

*  Huber,  Deutsch: Arch.  f.  klin.  Med.,  xlvii. 

6  Ramon  de  la  Sota  y  Lastra,  Rev.  vied,  de  Sevilla,  1883. 

■^  Photiades,  Int.  Zentralbl.  f.  Laryng.,  1884. 

^  Maissurianz,  St.  Petersb.  vied.  Wochenschr.,  1883. 

44 


700  THE   ANIMAL   PARASITES   OF   MAN 

Schmolitschew^  is  an  interesting  one ;  it  was  that  of  a  woman  who 
for  four  days  had  suffered  from  violent  haem^optysis,  the  cause  of 
which  was  a  leech  that  was  fixed  on  the  laryngeal  wall  of  the  epiglottis 
close  above  the  vocal  cords.  In  his  case  (that  of  a  soldier),  Godet^ 
was  forced  to  perform  thyrotomy  to  remove  the  leech  from  the  larynx. 
Ficano^  removed  a  live  leech  wdth  the  forceps  from  the  lower 
laryngeal  cavity  in  a  man,  aged  30.  Massei*  reports  a  similar  case. 
The  case  reported  by  Winternitz  and  Karbinski^  was  that  of  a 
peasant  girl,  aged  16,  who  suffered  from  coughing,  hoarseness,  and 
blood-stained  expectoration  ;  a  leech  had  lodged  on  the  root  of  the 
epiglottis.  Aubert^  removed  a  leech  from  the  larynx  of  a  woman 
after  the  performance  of  tracheotomy.  Seifert^  reports  three  cases  : 
in  the  first  the  leech  had  become  fixed  to  the  left  vocal  cord,  in  the 
second  it  was  found  in  the  lower  laryngeal  cavity,  and  in  the  third 
on  the  border  of  the  left  ligamentum  aryepiglotticum.  Leone^  has 
published  the  case  of  a  leech  in  the  larynx,  Martin^  two  cases  with 
the  leech  lodged  in  the  lower  laryngeal  cavity,  Be'rthoud^^  a  similar 
case,  Palazzolo^^  two  such  cases,  Panzat^^  one  case  (lower  laryngeal 
cavity).  Moucharinski^^  reports  a  case  in  which  the  leech  had  stayed 
more  than  twenty  days  in  the  larynx.  Martin^*  easily  removed  a 
leech  from  the  posterior  portion  of  the  vocal  cord  with  the  forceps. 
Vieus  and  Nepeon^^  record  a  case  of  a  leech  in  the  larynx.  It  is 
quite  exceptional  for  leeches  to  gain  access  to  the  trachea  ;  cases  of 
this  kind  have  been  recorded  by  Aubert,^^  Vicano,^^  Ridola^^  and 
Tapin^^  (the  leech  was  firmly  fixed  to  the  bifurcation  and  caused 
coughing,  haemoptysis  and  attacks  of  asphyxia  ;  it  was  easily  removed 
by  the  aid  of  a  tracheal  tube).  Now  and  then  leeches  are  found  in 
the  nose. 

Lusitanus^^  relates  the  case  of  a  man  who  suffered  from  severe 
headaches.  A  medical  man  ordered  the  application  of  a  leech  to  the 
anterior   portion  of  the    nostril.     Owing   to   the  carelessness  of   the 

'  Schmolitschew,  Wratsch,  1884. 
2  Godet,  Arch,  de  Med.  et  Pharm.  milit.,  1887. 

»  Ficano,  Rev.  de  Laryng.,  1890.  *  Massei,  Int.  Journ.  of  Laryng.,  1890. 

^  Winternitz  and  Karbinski,  Prag.  vied.  Wochenschr.^  1890. 
«  Aubert,  Echo  med.,  1891.  '^  Seifert,  Rev.  de  Laryng.,  1893. 

^  Leone,  Boll.  del.  mal.  delP  orecchio,  etc.,  1892. 
®  Martin,  Arch,  de  Med.  et  Pharm.  milit.,  1891. 

*o  Berlhoud,  ibid.,  1893.  "  Palazzolo,  Boll.  del.  mal.  delt  orecchio,  1895. 

'2  Panzat,  Arch,  de  Mid.  et  Pharm.  milit.,  1896. 
"  Moucharinski,  Wratsch,  1896. 
1^  Martin,  Rev.  barcelon  de  enf.  de  oido,  1906. 
'^  Vieus  and  Nepeon,  Monatsschr.  f.  Ohrenheilk.,  1884. 
*6  Aubert,  Echo  med.,  October  12,  1891. 
'■^  Vicano,  Boll.  del.  mal.  deW  orecchio,  etc.,  1892,  ix. 

'^  Ridola,  Arch.  ital.  di  Laryng.,  1894,  ii.  '^  Tapin,  Siglo  med.,  March  16,  1907. 

^  Lusitanus,  see  Seifert  in  Ileymann's  "  Handb.,"  p.  599. 


SUPPLEMENT 


701 


surgeon  the  leech  crawled  right  into  the  nose ;  it  was  impossible  to 
extract  the  leech  or  to  kill  it,  and  it  produced  a  severe  haemorrhage 
which  led  to  the  death  of  the  patient  within  two  days.  In  a  case  recorded 
by  Sinclair/  a  leech,  Ha'tnopsis  sangiiisuga,  gained  access  to  the  nose 
of  a  boy,  aged  3  ;  it  remained  there  a  fortnight ;  it  caused  frequent 
attacks  of  epistaxis  and  in  the  end  it  was  removed  by  means  of 
forceps.  Condorelli-Francaviglia^  records  a  case  in  which  severe 
epistaxis  was  caused  by  a  leech  which  had  probably  entered  the 
anterior  portion  of  the  left  nostril  by  way  of  the  pharynx  and  become 
tightly  fixed  there.  It  was  seen  by  posterior  rhinoscopy,  and 
was  removed  from  in  front  by  means  of  slightly  curved  forceps. 
Sota  y  Lastra^  mentions  the  occurrence  of  leeches  in  the  nose,  and 
Keng^  reports  the  case  of  nasal  obstruction  from  a  leech.  The 
removal  of  leeches  is  effected  by  means  of  injections  or  by  the  direct 
sprinkling  of  salt  or  acid  solutions  on  their  bodies,  which  brings  about 
their  detachment.  When  possible  a  previous  attempt  should  be  made 
to  seize  them  with  forceps  so  as  to  make  their  immediate  extraction 
possible.  The  species  of  Haemadipsa  (Looss*^)  live  in  tropical  regions 
in  moist  places  on  the  ground  or  in  the  jungle.  They  climb  bushes 
and  even  trees  with  astonishing  rapidity  upon  the  approach  of  larger 
animals  and  also  of  man  (whom  they  clearly  recognize  from  the 
vibration  of  the  ground  caused  by  footsteps).  From  thence  they  let 
themselves  fall  on  their  victims  to  suck  their  blood.  Their  bites  are 
generally  painless,  and  of  themselves  not  dangerous,  but  if  they  are 
unusually  numerous  they  rapidly  accumulate  on  the  body  in  large 
numbers  and  give  rise  to  marked  debility  and,  if  the  wound  become 
infected,  to  severe  complications  and  even  death.  On  the  other  hand, 
under  careful  treatment  the  wounds  heal  easily  and  fairly  rapidly. 

Firm  leather  and  firmly  adhering  clothes  afford  no  certain  pro- 
tection against  the  attacks  of  these  leeches,  as  they  know  how  to 
force  themselves  with  extraordinary  rapidity  through  the  narrowest 
interstices  betw^een  the  clothes  and  thus  gain  access  to  the  skin. 
When  they  have  sucked  their  fill — and  this  may  take  several  hours  to 
accomplish — they  fall  off  of  themselves.  To  effect  an  earlier  removal 
drops  of  irritative  or  corrosive  fluids  are  employed  (salt  solutions, 
acids,  etc.).  Tearing  away  the  leech  by  force  should  be  avoided, 
as  in  this  way  portions  of  the  leech's  body  may  be  left  behind  in  the 
wound  and  inflammation  be  set  up. 


1  Sinclair,  BriL  Med.  Journ.,  June  20,  1885,  i. 

■^  Condorelli- Franca viglia,  Spallangini,  1892. 

'^  Sota  y  Lastra,  Rev.  vied,  de  Sevilla,  1887. 

*  Keng,  Scot.  Med.  and  Surg.  Journ.^  October,  1899. 

«  Looss,  "  Handb.  d.  Tropenkrankh.,"  v.  Mense,  i,  p.  194. 


702  THE   ANIMAL   PARASITES   OF   MAN 


ARTHROPODA. 

Leptus  autumnalis  (Grass,  Harvest,  or  Gooseberry  Mite^). 

In  the  hot  season  of  the  year,  that  is,  during  the  months  of  July 
and   August,    it   is   noticed    that    those    people   who   stray   amongst 
syringa    bushes    or    who    pick    gooseberries    or    kidney   beans    are 
attacked  by  the  Leptus  autninnalis.     On  the  uncovered  parts  of  the 
body  there  appear  numerous  red  spots  and  papules,  which  itch  and 
burn  smartly.     The  itching  does  not  commence  diffusely,  as  in  the 
case    of    scabies    (MacLennars^),    but    is   Hmited    to    the    particular 
points  where  the  parasite  is  situated.     There  are  especial  outbreaks 
of  itching  in  the  morning,  arising  perhaps  from  the  hatching  of  ova 
in  the  host  after    lying   in   the  warmth    of    the  bed.^      Leptus   fre- 
quently provokes  general  erythema,  eczematization  or  severe  feverish 
urticaria,  which  in  France  is  known  by  the  narne  of  fievre  de  grain 
(Megnieu,   Besnier^).      If    the    individual    efflorescences  be  carefully 
examined,  there  will   be  noticed  almost  without  exception   a  minute 
boss  towards  the  centre,  noticeable  by  its  yellowish-red   colour.     If 
an  attempt  is  made  to  remove  it  w^ith  the  point  of  a  needle  or  to 
scrape  it  off  the  surface,  one  can  often  perceive,  even  with  the  naked 
eye,  a  small  reddish  creature  moving  actively  about.     The  treatment 
of  these   very  troublesome  symptoms   consists    in  warm   baths  with 
soapy   lavages,  also  lavages  with  alcohol,  spirit  salmiac  (G.  P.),  5  per 
cent,  carbol  or  creolin  solution,  diluted  vinegar,  benzine,  emulsions  of 
balsam  of  Peru,  rubbing  in  sulphur  ointment  (Sandwith'^)  ;  ointments 
of  creosote  or  eucalyptus  are  recommended.     Other  grass  and  grain 
mites  also  occasionally  penetrate  the  skin  of  man  and  produce  transi- 
tory  but   sometimes    very   severe    eruptions,    urticaria   and   eczema 
papulosum,  as  Geber^  and  subsequent  to  him  Josai'  have  reported 
of  the  barley  mite.     In  sensitive  individuals  the  skin  becomes  bright 
red,  to  a  greater  or  less  extent  their  temperature  is  raised   and   fre- 
quently  slight    febrile  affections   are    present.     If   the    inflammatory 

'  There  is  no  reason  for  calling  this  the  gooseberry  mite.  It  is  rarely  found  on  this 
fruit.     The  gooseberry  mite  is  Bryolia  pi-etiosa. 

-  MacLennars,  Lancet,  1905. 

'  [This  cannot  be  the  case,  as  Leptus  aututnnalis  is  the  larval  form  of  Trombidiiun  hola- 
sericeum. — F.  V.  T.] 

^  Sack,  "Handb.  d.  Hautkrankh.,"  v.  Mracek,  1907. 

*  Sand  with,  Lancet,  1905. 

^  Geber,  "  Ilandbuch  d.  Hautkrankh.,"  in  v.  Ziemssen's  "  Handbuch  d.  spez.  Pathol, 
u.  Therap.,"  1884,  xiv. 

^  Josai. 


SUPPLEMENT  703 

skin  symptoms  have  reached  their  culminating  point  after  three  or 
four  days  and  no  fresh  comphcations  arise,  they  only  remain  for 
a  short  while,  the  effects  of  scratching  and  pigment  spots  being 
left. 

Kedani,  Akaneesch   (The  Japanese  River  or  Inundation  Disease). 

This  disease  is  only  known  in  Japan,  and  is  limited  to  the 
neighbourhood  of  some  great  rivers  on  the  west  coast.  The  people 
mostly  attacked  are  those  who  cut  the  hemp  harvest  in  the  infected 
localities,  occasionally  those  who  transport  it  or  come  into  contact 
with  it  (Looss^).  The  disease  is  frequently  manifested  in  the  form 
of  indefinite  disturbances  of  the  general  condition ;  it  commences 
generally  on  the  sixth  day  after  the  presumed  infection  with  rigors, 
headaches,  feeling  of  weakness,  sw^elling  of  the  lymphatic  glands  in 
the  loin  or  in  the  arm-pits  ;  in  the  periphery  a  black  dry  scab  is 
formed.  In  addition  there  is  an  intense  conjunctivitis,  and  added 
to  symptoms  of  fever  an  exanthema  resembling  measles  that  lasts 
from  four  to  seven  days.  There  is  frequent  delirium  and  difHculty 
of  hearing  which  persist  for  a  long  while.  Obstinate  constipation  is 
a  striking  symptom.  At  the  end  of  a  fortnight,  earlier  in  slighter 
cases,  the  fever  commences  to  abate  and  a  rapid  convalescence  sets 
in.  In  pregnant  women  abortion  with  fatal  issue  is  frequent.  With 
regard  to  prophylaxis,  Baelz^  recommends  as  rapid  a  cultivation  of 
the  soil  as  possible,  which  has  led  to  a  speedy  disappearance  of 
the  disease .  in  districts  where  it  was  once  dreaded.  Treatment  is 
symptomatic.  Japanese  do  not  tolerate  antipyretic  drugs  as  well 
as  Europeans. 

Dermanyssus  gallinae  (avium). 

During  the  day  the  resort  of  bird  mites  is  in  the  droppings  and 
in  the  woodwork,  etc.,  of  cages  in  which  canaries,  crossbills  and 
parrots  are  kept ;  in  the  crevices  of  doors,  in  the  chinks  between 
the  board  planks  of  bedsteads,  so  that  at  night  they  may  seek  some 
domestic  animal  to  suck  the  blood  and  so  satisfy  their  hunger.  It 
is  by  no  means  rare  for  young  animals,  chickens  and  unfledged 
pigeons,  etc.,  to  perish  in  consequence  of  the  great  loss  of  blood. 
This  nocturnal  habit  of  life  explains  why  no  mites  can  be  found 
during  the  day  in  spite  of  the  most  careful  examination  of  the  human 
body,  to  which  they  may  be  transmitted.  On  the  uncovered  parts 
of  the  body  they  not  only  cause  severe  irritation,  but  also  severe 
diffuse  itching  erythema  and  eczema.     Thorough  disinfection  of  the 

'   Looss,  '•  Handbuch  d.  Tropenkrankh.,"  v.  Mense,  p.  195. 
-  Baelz,  Virchow's  Archiv,  Ixxviii. 


704 


THE   ANIMAL   PARASITES   OF   MAN 


cages  by  hot  solution  of  caustic  potash,  in  addition,  sprinkHng  over 
with  tar,  red  carbohc  acid  or  petroleum,  thoroughly  powdering  over 
the  birds  with  flores  pyrethrae,  washing  with  water  containing  oleum 
anisi,  washing  the  walls,  doors  and  bedsteads  with  soap,  disinfection 
of  the  mattresses,  linen  and  clothes,  will  protect  against  further 
infection.  In  the  case  of  man  the  disease  needs  no  special  treatment, 
as  the  eruptions  generally  disappear  after  some  days.  Heinecke^ 
recommends  lavages  with  i  per  cent,  carbolic  acid  solution.  \_Vide 
also  p.  492  in  body  of  this  work. — F.  V.  T.] 

[Dermanyssus  hirundinis,  Hermann,  is  identical  with  this  species. 
By  far  the  best  treatment  is  with  paraffin  or  kerosene  oil  applied  to 
the  places  where  they  pass  the  day. — F.  V.  T.] 

Ixodes  reduvius  (ricinus). 

The  female  is  occasionally  transmitted  to  the  human  skin,  and 
bores  its  proboscis  deep  into  it  and  sucks  itself  full  of  blood.  At 
sensitive  points  of  the  cutaneous  surface — for  example  over  the  skin 
of  the  penis — a  feeling  of  severe  pain  is  produced.  Buy's^  observations 
as  to  the  geographical  distribution  of  the  Ixodina  show  that  in  all 
lands  in  which  cattle,  horses,  sheep  and  dogs  exist,  Ixodince  are  to  be 
found.  Recent  observations  show  that  the  Ixodince  play  an  important 
part  in  the  transmission  of  Hsemosporidia  {vide  body  of  work, 
pp.  493,  494).  Sprinkling  with  oil,  vaseline,  benzine,  ether,  petroleum, 
naphtha,  turpentine  (Jelgenum^),  will  easily  lead  to  the  removal  of 
the  parasite ;  if  the  body  is  torn  away  with  violence  and  the  proboscis 
is  left  sticking  in  the  skin,  the  presence  of  the  latter  will  give  rise  to 
inflammation  and  suppuration. 

Sarcoptes  scabiei  (Scabies). 

The  disease  produced  by  Sarcoptes  scabiei  shows  itself  in  poly- 
morphous areas,  such  as  accompany  eczema,  and  are  produced  on 
the  one  hand  by  the  Sarcoptes  alone  and  on  the  other  hand  by 
the  scratching  with  the  nails.  The  localization  of  both  kinds  of 
efflorescences  is  different  from  those  which  are  produced  by  the 
Sarcoptes  ;  they  occur  as  papules,  vesicles,  pustules  and  mite-tracks, 
and  their  usual  situation  is  between  the  fingers,  on  the  ulnar  border 
of  the  hand,  on  the  wrist,  on  the  palm  of  the  hand,  on  the  anterior 
border  of  the  axilla,  on  the  penis  and  at  the  base  of  the  thorax.  The 
excoriations  are  situated  on  the  forearm,  over  the  thigh,  over  the 
abdomen,  and  may  be  distributed  in  greater  or  less  degree  over  the 

'  Heinecke,  Milnch.  med.  Wochenschr.,  1901. 

-  Buy,  "  Histoire  naturelle  et  medicale  des  Ixodes,"   "These  de  Lyon,"  1906. 

^  Jelgenum,  Med.  Weekbladv.  Noord-  en  Ztiid-Nederland,  1901,  i,  No.  24. 


SUPPLEMENT  705 

whole  body  ;  the  back  and  the  face  only  remain  free.  The  symptoms 
consist  in  violent  itching,  the  onset  of  which  specially  takes  place 
at  night. 

The  mite-tracks  are  fine  curving  lines,  curved  like  a,  u,  c,  or  s, 
which  appear  as  if  they  had  been  scratched  with  a  fine  needle.  Upon 
closer  examination  with  the  magnifying  glass  one  sees  in  their  course 
sm^ll  openings.  These  openings,  in  persons  who  keep  themselves 
clean,  are  scarcely  coloured ;  but  in  patients  whose  occupations 
necessitate  their  being  associated  with  coloured  or  dirty  substances, 
they  are  dark.  The  length  of  the  tracks  varies  from  some  millimetres  to 
1 J  to  2  cm.  They  are  at  the  one  end,  where  the  Sarcoptes  is  embedded 
in  the  epidermis,  widened  like  a  funnel  and  slightly  exfoliated.  The 
track  at  this  point  is  sharply  defined  ;  the  mite  shows  through  the 
epidermis  as  a  yellowish  round  point.  In  the  course  of  the  track  there 
develop  papulae,  vesicles  or  pustules,  which  raise  the  level  of  the  track. 
The  intensity  of  these  inflammatory  appearances  depends  upon  the 
susceptibility  of  the  human  individual  and  upon  the  capability  of 
the  reaction  of  the  skin.  There  are  people  in  whom  scarcely 
any  inflammatory  symptoms  make  their  appearance  ;  on  the  other 
hand  there  are  some,  especially  children  and  lymphatic  individuals^ 
in  whom  severe  impetiginous  ecthymatous  pustules,  together  with 
their  sequelae,  are  set  up. 

The  results  produced  by  scratching  consist  in  papules,  which 
usually  bear  a  small  scab  of  blood,  and  are  arranged  in  the  form  of 
striae,  in  eczematous  surfaces,  weeping  or  sanguineous  scabs,  vesicles, 
pustules,  etc.  The  complications  that  set  in  are  frequently  urticaria 
and  even  furuncles,  lymphangitis  and  inflammation  of  the  glands, 
which  now  and  then  is  followed  by  the  formation  of  abscesses  in  the 
glands. 

The  duration  of  the  disease  is  unlimited  ;  when  untreated  it  leads 
to  a  form  of  rare  occurrence,  that  of  scabies  norvegica^ ;  in  this  the 
collection  of  crusts  and  scales,  in  which  a  quantity  of  dead  mites, 
larvae  and  ova  are  present,  may  become  colossal. 

The  symptoms  of  scabies  abate  in  the  presence  of  intercurrent 
acute  diseases  and  reappear  after  the  malady  is  over.  The  fact  has 
for  long  contributed  to  the  idea  of  scabies  being  regarded  as  a  disease 
capable  of  being  ^*  driven  in"  upon  the  internal  organs  and  forming 
metastases. 

The  diagnosis  is  rendered  certain  upon  the  discovery  of  a  track. 
Traces  of  scratching  on  the  extremities  and  on  the  abdomen,  papular 
or  pustular  efflorescences  between  the  fingers,  toes,  in  the  neighbour- 
hood of  the  wrist,  of  the  elbow,  on  the  anterior  border  of  the  arm-pit, 
on  the  tuber  ischii,  in  the  girdle  region,  and  especially  the  presence  of 

*  [This  is  produced  by  a  distinct  species,  vide  pp.  519-20. — F.  V.  T.] 


706  THE   ANIMAL   PARASITES    OF   MAN 

disintegrated  tracts  over  the  penis  (prepuce  and  glans),  will  allow  of 
the  diagnosis  being  made.  Certain  occupational  eczemas  (grocers, 
lime- workers,  maltsters,  bakers  and  others),  also  prurigo,  must  be 
borne  in  mind  when  diagnosing  this  disease. 

The  prognosis  is  always  a  favourable  one.  Even  after  such  a 
long  duration  and  after  such  severe  symptoms  the  disease  may 
completely  clear  up.  There  are,  however,  frequently  left  behind  post- 
scabious  inflammatous  and  pruriginous  conditions  which  only  yield 
after  protracted  treatment.  Scabiophilia,  which  persists  in  certain 
patients  for  a  long  time  after  the  scabies  has  been  cured,  must  here 
be  mentioned. 

In  the  treatment  of  scabies  four  points  must  be  kept  in  view, 
(i)  The  mites  and  the  ova  must  be  killed  by  the  treatment ;  (2)  the 
treatment  must  have  regard  to  the  intensity  of  the  inflammatory 
symptoms ;  (3)  the  clothes  (body-linen)  of  the  patients  must  be 
disinfected  ;  the  bed-linen,  the  beds  and  the  bedsteads  must  be 
cleansed  ;  (4)  when  a  person  suffers  from  scabies  his  entourage 
must  be  examined,  and  all  diseased  conditions  treated  in  the  same 
way  as  under  (3). 

The  treatment  (i)  should  be  preceded  by  a  bath  wath  thorough 

soap   ablution,  and  when   the  inflammatory  symptoms   are   not  too 

severe,  with  green   soap.     After   the  bath  the  skin  is  dried  and  the 

scabies   remedy    proper   applied    in    warmth.      Sulphur   preparations 

receive  first  consideration ;  among  such  Vlemingkz's  mixture  occupies 

a  prominent  position  ;  this  is  rubbed  in  for  half  an  hour  by  means  of 

a   strong   camel-hair    brush,    to    be    followed    by   another    bath    and 

powder  applications  after  drying.     Repeat  this  method  for  three  days 

one  after  the  other,  or  for  two  days,  and  a  third  time  eight  days  later. 

The  latter  method  is  worthy  of  recommendation  as  the  ov^a,  which 

perhaps  resist  the  parasiticide  action,  have  by  this  time  developed  into 

larvae,  and  the   latter   can    then    be    destroyed  with    certainty.     The 

remaining  sulphur  preparations,  which  are  specially  employed  in  the 

form  of  ointments,  are  more  complex,  as  the  ointment  should  remain 

on  the  skin.     Helmerisch's  and  Wilkinson's  ointments  are  the  kinds 

specially  employed.     Nagelschmidt^  recommends  thiopinol  as  a  very 

suitable  sulphur   preparation    in    the   form   of   baths   or  as  a   10   or 

5  per  cent,  ointment  in  the  following  way  :  Upon  his  reception  the 

patient  is  given    a    thiopinol   bath,  in   which   he  remains    for   thirty 

minutes.     Immediately  afterwards  30  to  40  grm.  10  per  cent,  thiopinol 

vaseline  is  carefully  rubbed  in.     The  rubbing  is  repeated  daily,  and 

the  treatment  is  concluded  on  the  second  to  fourth  day  with  a  second 

thiopinol   bath.     Thiopinol    produces    no    more    irritation    than    the 

ordinary  sulphur  ointments  ;  it  is,  however,  much  more  penetrative 

and  more  capable  of  absorption. 

'  Nagelschmidt,  Med.  Klin.,  1907,  xxxv. 


SUPPLEMENT  707 

We  frequently  make  use  of  Kaposi's  naphthol  ointment,  as  it 
renders  the  skin  supple,  causes  proportionately  little  irritation,  and 
has  but  little  smell.  Treatment  with  balsam  of  Peru  is  certainly 
expensive,  but  in  the  slighter  attacks  it  is  relatively  the  simplest.  We 
give  the  patient  a  bath,  have  him  thoroughly  dried  and  rub  in  30  to 
40  to  50  grm.  balsam  of  Peru  carefully  and  evenly  all  over,  wrap 
him  in  a  covering  of  wool,  and  make  him  rest  in  bed  for  twelve  to 
fifteen  hours,  to  be  followed  by  a  bath  with  careful  cleansing  with 
soap ;  this  treatment  need  rarely  be  repeated.  The  balsam  of  Peru 
can  be  applied  undiluted  for  the  rubbings  or  mixed  with  ung. 
glycerini,  or  resorbin  or  glycerine  in  equal  parts.  [Norman  Walker 
uses  balsam  of  Peru  J  oz.  dissolved  in  rectified  spirit ;  to  be  painted 
on  with  a  brush.] — ].  P.  S.  The  manufacturers  name  the  undiluted 
product  of  the  active  constituent  of  balsam  of  Peru,  benzoic  acid 
benzyl-ester,  Peruscabin.  For  the  treatment  of  scabies  it  is  recom- 
mended by  Sachs^  that  it  should  only  be  administered  when  mixed 
with  ricinus  oil,  under  the  name  of  Peru  oil,  in  applications  repeated 
three  times  within  thirty-six  hours. 

Sack"2  also  considers  Peru  oil  a  non-irritant,  effectual,  pleasant, 
inodorous  and  non-staining  drug.  But  he  only  allows  the  applica- 
tions to  be  used  every  twelve  hours  for  three  to  four  consecutive  days 
(altogether  200  to  300  grm.  of  Peru  oil  are  requisite),  and  after  the 
sixth  or  seventh  rubbing  a  bath  should  be  taken  with  the  use  of 
Dutch  soap.  Juliusberg^  considers  this  treatment  specially  suited 
for  private  practice.  Another  modern  drug  is  epicarin  (/3-oxy- 
naphthyl-ortho-oxy-meta-tolyol  acid)  ;  this  is  applied  in  10  to  20  per 
cent,  ointments  (Pfeiffenberger^),  epicarin  7*0  grm.,  cretae  alb.  2*0  grm., 
vasel.  flavi  30*0  grm.,  lanolin  i5"o  grm.,  axungia  poric.  45*0  grm. 
(Rille^) ;  epicarin  15-0  grm.,  sapon.  virid.  5*0  grm.,  axung.  poric.  100*0 
grm.,  cretae  alb.  ico  grm.  (Kraus^)  ;  for  children,  epicarin  5*0  grm., 
lanolin  90*0  grm.,  ol.  olivar.  lo'o  grm.  (Kaposi').  Siebert^  lays  stress 
upon  the  odourlessness  and  colourlessness  of  epicarm  ointment  as  a 
strong  reason  for  its  use,  and  points  out  that  it  is  a  harmless  drug,  the 
action  of  which  is  certain.  Endermol  (salicylic  acid  ointment)  has  a 
destructive  action  on  the  mites  even  in  a  O'X  per  cent,  ointment 
(Wolters,^  Demitsch^^)  ;  it  is,  however,  very  expensive  and  not  wholly 
free  from  danger ;  and  the  same  applies  to  nicotiana  soap  (Taenzer,^' 
Schumann^^j. 

'  Sachs,  Deutsche  med.  Wochenschr.^  1900. 

'^  Sack,     "  Handb.  d.    Hautkrankh.,"  v.  Mracek. 

'^  Juliusberg,  Therap.  Monatsh.,  1901. 

^  Pfeiffenberger,  Klin,  therap.  Wochenschr.,   1900.         ''  Rille,  "  Die  Heilkunde,"  1900. 

^  Kraus,  Allg.  wien.  med.  Zeit.^  1900.  '^  Kaposi,  Wien.  nied.  Wochenschr.^  1 900. 

•*  Siebert,  Munch,  med.  Wochenschr.^  1900.  '-^  Wolters,  Therap.  Monatsh.,  1898. 

'°  Demitsch,  Wratsch,  1905,  iv.  "  Taenzer,  Monatsh.  f.  prakt.  Derm.,  xxi. 

^'^  Schumann,  Allg.  med.  Central-Zeitg.,  1901. 


yoS  THE   ANIMAL   PARASITES   OF   MAN 

To  give  an  account  in  detail  of  the  drugs  and  methods— old  and 
new — used  in  the  treatment  of  scabies  would  far  outrun  the  limits  of 
this  work. 

Demodex  folliculorum. 

It  is  not  yet  certain  whether  the  Demodex  folliculorum  is  capable 
of  developing  pathological  conditions  in  man.  VeieP  assumes  that 
the  hair  follicle  mite  has  no  connection  either  with  the  formation 
of  comedones  or  even  with  sebaceous  gland  disease.  Kaposi^  con- 
siders that  they  cause  no  disease  in  man  and  cannot  be  regarded  as 
a  cause  of  acne.  Saalfeld^  clearly  adheres  to  the  same  standpoint, 
similarly  so  Jessner,*  who,  when  discussing  comedones,  makes  no 
mention  of  acne  of  hair  follicle  mites.  WeyF  and  Geber^  adhere  to 
the  opinion  that  the  presence  of  a  Demodex  in  man  in  contradis- 
tinction to  its  presence  in  animals  possesses  absolutely  no  pathogenic 
influence.  On  the  other  hand  de  Amicis,'  Majochi,^  and  Dubreuilh'^ 
report  single  cases  of  pronounced  circumscribed  clear  brown 
pigmentations  which  they  attribute  to  Demodex  follicidorum.  In  all 
these  cases,  moreover,  as  regards  localization  the  affection  had  a 
certain  resemblance  to  pityriasis  versicolor  ;  nevertheless,  in  the  scales 
separated  off  with  the  scalpel  no  fungi  were  found,  but  on  the  other 
hand  Demodices  in  moderate  quantity.  In  his  earlier  cases  Majochi 
has  seen  the  Demodex  in  the  secretion  from  meibomian  glands  and 
had  claimed  it  to  be  the  excitant  of  chalazion  and,  as  Mibelli^^  did,, 
considered  it  to  be  the  cause  of  some  diseases  of  the  eyelids.  Ivers^^ 
found  the  parasite  in  69  per  cent,  of  normal  borders  of  the  eyelids, 
and  attributes  a  pathological  signification  to  it.  Hiinsche^^  and 
Mulder^^  arrive  at  the  same  conclusions  ;  in  the  light  of  their  investiga- 
tions the  Demodex  is  found  as  a  constant  accessory — certainly  not  in 
the  meibomian  glands,  as  it  is  limited  only  to  the  internal  part  of  the 
hair  follicle.  Lewandowsky^*  considers  that  it  can  hardly  be  demon- 
strated at  present  that  the  same  parasite  which  in  individual  specimens 
causes  no  symptoms  is  capable  of  producing  pathological  conditions 
when  markedly  increased  in  numbers. 

Treatment  is  by  the  removal  of  the  comedones,  above  all,  by  their 
mechanical   removal   by    pressure   with    a   watch-key   and   with   the 

1  Veiel,  V.  Ziemssen's  "  Handb.  d.  spez.  Path.  u.  Therap.,"  1884,  xiv. 

2  Kaposi,  "  Path.  u.  Therap.  d.  Hautkrankh.,"  1899. 

3  Saalfeld,  Lesser's  "  Encyclop.  d.  Haut-  u.  Geschlechtskrankh.,"  1900. 
*  Jessner,  *' Kompend.  d.  Hautkrankh.,"  1906,  3rd  ed. 

*,  ^  Weyl  and  Geber,  v.  Ziemssen's  "  Handb.  d.  spez.  Path.  u.  Therap.,"  1884,  xiv. 

■^  de  Amicis,  quoted  by  Lewandowsky.  *^  Majochi,  Centralbl.f.  Bakt.,  xxv. 

^  Dubreuilh,  La  Prat.  Derm.,  Paris,  1901. 
'°  Mibelli,  quoted  by  Lewandowsky.  "  Ivers,  ibid. 

'■^  Hiinsche,  MiXnch.  fried.  Wochenschr.,  1900,  xiv. 
'3  Mulder,  Weekbl.  v.  het  Nederl.  Itjdschr.  v.  Geneesk.,  1889. 
'*  Lewandowsky,  Deutsch.  med.  Wochenschr. ^  1907,  xx. 


SUPPLEMENT  709 

various  comedo-compressors,  and  by  subsequent  cleansing  of  the 
skin  with  ether,  benzine  or  spirit.  If  the  eyehds  should  be  affected 
with  blepharitis  due  to  the  presence  of  Demodex  in  large  numbers, 
epilation  and  administration  of  a  parasiticide  is  recommended. 

Dennodex  folliculorunn  canis. 

Transmission  from  dog  to  man  is  in  any  case  very  rare,  and  by 
man}^  its  occurrence  is  generally  doubted.  Nevertheless  Gruby^  and 
Remak^  claim  that  it  is  transmissible — an  opinion  which  has  also 
been  shared  by  Neumann^  and  Ziirn.*  The  latter  saw  in  the  case 
of  a  married  couple  who  had  the  care  of  mangy  dogs  the  onset  of 
diseased  areas  on  their  hands  and  feet,  which  were  like  those  on  the 
dogs  and  contained  the  same  parasites. 

A.  Babes^  also  reports  several  observations  which  go  to  show 
that  persons  who,  to  some  extent,  have  been  shown  to  have  been  in 
contact  with  mange-stricken  dogs  have  been  attacked  by  a  scabies- 
like eruption  localized  over  the  thorax,  abdomen,  back  and  extremi- 
ties ;  large  numbers  of  Demodices  were  found  in  the  follicular 
pustules.  Lewandowsky^  reports  one  case — that  of  an  Italian 
workman,  who  suffered  from  an  outbreak  on  the  face,  like  impetigo  ; 
there  was  crust  formation  and  at  the  edge  of  the  crusts  the  epidermis 
appeared  like  a  narrow  row  or  border  of  vesicles.  A  small  portion 
of  the  covering  of  the  row  of  vesicles  was  lifted  off,  and  after  slight 
warming  examined  in  40  per  cent,  liquor  potassae.  In  this  a  large 
number  of  animal  parasites  of  the  Demodex  group  were  found,  and 
without  doubt  Demodex  folliciiloruni  canis  alone.  Hiinsche^  assumes 
that  Demodex  folliculorum  penetrates  into  the  tissues  and  produces 
abscesses. 

Treatment  first  consisted  in  dusting  with  zinc  amyl  powder,  but 
after  four  days  there  was  no  change.  After  the  regular  use  of  xero- 
form  as  a  powder  application,  the  affection  cleared  up  within 
fourteen  days. 

INSECTA. 

Pediculus  capitis  (Pediculus  capitis)  (Head  Louse). 

We  find  Pediculus  capitis  in  very  young  children  and  in  others 
more  growm  up  to  be  the  incessant  and  frequent  cause  of  impeti- 
ginous crust-forming  eczemas.     It  is  more  frequent  in  girls  than  in 


'  Gruby,  quoted  by  Lewandowsky.  ^  Remak,  idid. 

^  Neumann,  z'di'd.  "*  Ziirn,  idtd. 

•'  Babes,  ibid. 

°  Lewandowsky,  Deutsch.  med.   Wachenschr.^  ^907,  xx. 

■^  Hunsche,  Milnch.  med.  JVochenschr.,  1900,  xlv. 


7IO  THE   ANIMAL   PARASITES   OF   MAN 

boys.  In  families  it  is  endemic,  in  schools  epidemic,  but  it  also 
occurs  in  fair  frequency  in  female  adults  (servant  maids,  waitresses) 
who  may  pay  little  attention  to  bodily  cleanliness.  The  puncture 
of  the  parasites  sets  up  a  severe  irritation,  which  leads  to  violent 
scratching.  The  consequences  of  this  are  the  formation  of  nodules 
and  pustules,  crusts  and  "weeping"  patches;  the  hairs  become  felted 
and  the  final  clinical  picture  is  that  of  plica  polonica.  The  conditions 
of  irritation  which  are  produced  by  these  parasites  and  then  by  the 
scratchings  of  the  impetiginous,  and  frequently  the  very  severe 
suppurative  processes  of  the  hair-bed,  lead  to  swellings  in  the  neck 
and  sometimes  even  to  glandular  suppurations.  The  eczematous 
processes  not  infrequently  extend  over  the  face,  the  neck  and  the 
thorax.  Blepharitis  and  conjunctivitis  may  be  due  to  Pediculns 
capitis. 

The  means  of  infection  are  often  very  remarkable.  Transmission 
from  one  individual  to  another  certainly  often  occurs,  but  infection 
may  take  place  in  railway  carriages  and  in  other  ways.  A  case  under 
the  observation  of  a  colleague  in  Frankfort  is  a  most  remarkable  one  : 
he  diagnosed  pediculosis  as  the  cause  of  a  head  eczema  occurring 
among  the  children  of  one  of  the  best  families  there.  The  infection 
took  place  through  dolls  adorned  with  human  hair,  in  which  the 
presence  of  nits  could  be  demonstrated. 

The  diagnosis  of  Pediculns  capitis  is  not  difficult  to  make  when 
the  hairs  and  hairy  scalp  are  carefully  examined  for  nits  and  living 
parasites.  In  better  families  it  is  a  good  plan  to  point  out  the 
corpora  delicti  to  their  possessors  and  to  make  them  aware  of  the 
possible  sources  of  infection. 

As  regards  treatment,  lotions  of  sabadill  vinegar  are  recom- 
mended ;  in  slighter  cases  these  are]  quite  sufficient.  In  severe  cases 
cure  will  not  result  unless  dressings  of  petroleum,  naphthol  ointment 
(5  to  10  per  cent.)  and  balsam  of  Peru  be  applied.  In  the  case  of 
plica  polonica,  the  hair  must  be  cut  quite  short  (even  in  adults)  so 
as  to  control  matting  of  the  hair.  To  get  rid  of  nits  from  hair  that 
is  not  matted,  careful  combing  and  washing  with  strongly  alkaline 
fluids  or  with  hot  vinegar  is  suitable. 


Pedlculus  vestimenti  (Clothes  Louse). 

The  clothes  louse  attacks  adults  by  preference,  and  with  especial 
frequency  old  and  emaciated  persons.  It  lives  in  the  clothes,  but 
derives  its  nourishment  from  the  body.  At  the  moment  at  which 
the  clothes  louse  inserts  its  proboscis  into  the  skin  the  person  experi- 
ences a  slight  sting,  which,  however,  at  once  ceases  to  hurt.  If  the 
body  of  the  louse  is  sucked  full  of  blood  it  falls  off  and  the  individual 


SUPPLEMENT  711 

has  rest  from  it  for  a  time.  A  wheal  develops  around  the  haemor- 
rhagic  area  of  the  bitten  spot  and  itches  severely.  The  itching  goes 
on  until  the  eruption  is  scratched  all  over.  This  is  followed  by  crust 
formation.  When  many  parasites  are  present  the  itching  reflexes 
become  more  severe,  and  the  patients  scratch  themselves  considerably 
and  make  long  marks  at  those  places  where  the  Pediculi  have  been. 
The  localization  of  the  scratching  effects  is  characteristic,  correspond- 
ing with  folds  between  portions  of  clothing  (regions  between  the 
shoulder-blades,  wrist  and  neck).  If  the  condition  lasts  for  a  month, 
the  scratching  effects  extend  over  the  whole  body,  and  secondary 
efflorescences  become  associated  with  it,  such  as  pustules,  ulcers  and 
eczemas.  Intermediate  between  this  we  find  cicatrices  and  pigmenta- 
tion, the  latter  under  certain  circumstances  extending  over  the  whole 
body.  Sulla,  Herod,  Cardinal  Dupet,  Philip  II,  and  others  are  said 
to  have  died  from  louse  disease.  That  even  at  present  many  human 
beings  are  exposed  to  the  danger  of  being  devoured  by  lice  is  a  fact 
that  we  have  had  the  opportunity  of  observing  on  several  occasions. 
Only  to  record  one  instance,  a  man,  aged  65,  was  received  into  our 
clinic  some  time  ago  in  an  absolutely  neglected  condition  (he  had 
been  slaying  for  some  weeks  in  a  stable,  lying  on  a  wretched  bed). 
The  whole  of  the  surface  of  his  body  was  covered  with  countless 
furuncles,  of  greater  and  less  size,  which  had  partly  become  changed 
into  undermined  ulcers.  Over  the  ulcers  and  beneath  their  undermined 
edges  Pediculi  were  swarming. 


Phthirius  ingulnalis  {Pediculus  pubis)  (Crab  Louse). 

The  transmission  of  these  parasites  generally  takes  place  during 
coitus,  and  therefore  they  especially  occur  in  the  pubes.  It  is  possible 
also  that  transmission  is  effected  through  dirty  clothes  and  bed-linen 
and  privy  seats. ^  Starting  from  the  pubes  the  animals  crawl  out  over 
the  other  parts  of  the  body  provided  with  hairs  to  the  abdominal 
wall  and  the  thorax  (so  far  as  these  parts  are  furnished  with  thick 
hair)  to  the  arm-pits,  the  beard,  the  eyebrows  ;  not,  however,  to  the 
hair  of  the  head,  or  rarely  so ;  among  our  numerous  cases  we  have 
never  met  with  an  example  of  the  crab  louse  attackmg  the  hair  of 
the  head. 

The  irritation  produced  by  the  crab  louse  is  extraordinarily  severe, 
especially  during  the  night,  as  the  warmth  of  the  bed  incites  the  lice 
to  active  sucking.     In  consequence  of  the  violent  scratching  indulged 


*  [A  case  of  infection  through  a  dirty  station  privy  in  Switzerland  came  to  my  knowledge 
in  1899,  and  numbers  of  pediculi  were  found  there. — F.  V.  T.] 


712  THE   ANIMAL   PARASITES   OF   MAN 

in,  eczemas  are  set  up  at  the  points  attacked,  and  these  often  spread 
to  the  neighbouring  parts  not  covered  with  hair. 

Of  special  interest  is  the  onset  of  maculcC  caeruleae  (taches  bleues) 
in  some  persons  affected  with  crab  Hce  (people  disposed  to  sweating 
seem  to  be  peculiarly  liable  to  these).  They  consist  in  pale  blue 
patches  of  various  size  and  shape,  varying  from  that  of  a  hemp-seed 
to  that  of  a  lentil,  and  again  to  that  of  a  nail  in  size  and  form.  These 
are  found  over  the  cutaneous  surface  of  the  abdomen,  thorax  and 
thigh,  and  are  often  only  seen  by  a  good  lateral  illumination.  Duguet^ 
considers  that  the  condition  is  a  toxic  erythema,  that  it  is  set  up,  on 
the  occasion  of  the  bite  of  the  parasite  penetrating  the  skin,  by  the 
poisonous  substance  derived  from  it.  Oppenheim^  considers  that  it 
is  a  colouring  substance  that  is  formed  in  the  salivary  glands  of  the 
parasites,  and  which  penetrates  the  skin  when  the  insects  bite,  and 
thus  forms  the  maculae  caeruleae.  We  have  on  several  occasions 
emulated  the  experiment  of  Duguet  (trituration  in  a  mortar  of  crab 
lice  freshly  taken  from  the  human  body  and  inoculating  the  mass 
thus  obtained  beneath  the  skin),  and  have  similarly  been  enabled  to 
produce  the  maculae  caeruleae  experimentally,  but  we  have  certainly 
been  unable  to  determine  which  of  the  hypotheses  is  the  correct 
one,  the  toxic  erythema  or  the  colouring  substance  inhibition 
theory. 

The  diagnosis  of  phthirasis  is  very  easy,  for  either  the  sexually 
mature  parasites  or  the  nits  are  found  on  the  hairs. 

As  regards  treatment,  grey  ointment  is  regarded  as  a  generally 
useful  application  ;  it  gives  rise,  however,  to  a  slight  eczema  of  the 
genitals,  espejially  in  males,  when  injudiciously  used.  Geber^ 
recommends  petroleum  or  balsam  of  Peru,  Oppenheim*  a  i  per  cent, 
sublimate  solution  for  lotions,  or  a  mixture  of  equal  parts  of  petroleum 
and  benzine  when  the  sublimate  cannot  be  borne.  The  use  of 
a  5  per  cent,  ointment  with  hydrarg.  oxid.  flavum  is  worth  consider- 
ing in  treatment  of  pediculosis  of  the  eyebrows  and  eyelashes.  The 
simplest  method  of  treatment,  and  one  with  a  radical  effect,  is  that  by 
sulphuric  ether  recommended  by  Thomer.^  It  certainly  produces 
a  sharp  burning  sensation,  but  the  living  parasites  and  nits  are 
destroyed  in  one  sitting.  We  prefer  ether  lotions  as  a  rule,  and  we 
thoroughly  rub  the  affected  parts  with  a  pad  of  wadding  well  soaked 
with  the  ether.     The  dead  parasites  and  the  nits  fall  on  to  what  lies 


*  Duguet,  Annal.  de  Derm.,  II  Ser.,  i. 

-  Oppenheim,  "Handb.  d.  Hauikrankh.,"  v.  Mracek,  1907. 

^  Geber,  see  Seifert,  Lesser's  "  Encyclop.,"  p.  387. 

^  Oppenheim,  loc.  cit. 

^  Thomer,  see  Seifert,  Lesser's  "  Encyclop.,"  p.  387. 


SUPPLEMENT  713 

beneath  when  the  rubbing  is  done  thoroughly,  and  the  burning  sen- 
sation caused  by  the  ether  only  lasts  a  few  minutes. 


Cimex  (Acanthia)  lectularia^  {Ciinex  lectiilariiis)  (Bed  Bug). 


The  puncture  in  the  skin  made  by  the  bed  bug  gives  rise  to  an 
extraordinary  amount  of  severe  itching  and  a  burning  sensation,  and 
Avhen  the  skin  is  sensitive  wheals  of  remarkable  size  {urticaria  ex 
cimicibus).  These  eruptions  that  cause  such  severe  itching  are 
scratched  by  those  attacked,  till  very  soon  blood  begins  to  flow,  and 
this  generally  leads  to  the  formation  of  a  dried  crust  of  blood  at  the 
point  of  eruption. 

The  diagnosis  is  not  always  easy,  as  urticaria  arising  in  other  ways 
frequently  leads  to  similar  vigorous  scratching  and  formation  of  crusts 
of  dried  blood.  Men  who  have  some  experience  in  this  matter  (for 
example,  commercial  travellers),  when  they  are  attacked  by  severe 
itching  at  night,  are  in  the  habit  of  striking  a  light  and  searching  in 
their  bed  and  body-linen  for  the  bugs,  in  order  to  be  able  to  hand 
over  the  corpora  delicti  to  the  landlord  if  need  be.  The  assumption 
that  the  bugs  in  the  East  play  an  actual  part  in  the  propagation  of 
tuberculosis  and  bubonic  plague  has  been  proved  by  investigations 
made  by  NuttalF  to  be  at  least  very  exaggerated  if  not  wholly  without 
foundation.  Further  investigations  may  decide  how  far  the  bugs 
participate  in  the  transmission  of  kala-azar,  as  is  believed  by  Rogers 
to  take  place. 

The  bed  bugs  must  be  exterminated  by  spraying  the  chinks  and 
joints  in  the  boards  with  petroleum  and  benzine,  pulling  up  the 
carpets  and  cleansing  the  bedsteads.  For  the  treatment  of  the  bite 
itself  the  methods  recommended  as  an  antidote  against  insects'  stings 
in  general  are  suitable  :  2  per  cent,  carbol  vaseline  (Rosen bach^), 
thymol  dissolved  in  spirit  (i  in  50^),  aethrol  or  deci-aethrol,  form- 
cethrol  (manufactured  by  Dr.  Nordlinger,  Florsheim  a.  /M.),  formoP 
(formol  15  parts,  xylol  5  parts,  acetone  44  parts,  Canada  balsam  i  part), 
with  the  aid  of  a  pad  of  wadding  placed  over  the  part  bitten,  lavages 
with  vinegar,  citron  juice  and  spirit  of  salmiac. 


1  Vide  genus  Cimex,  p.  534. 

■^  Nuttall,  see  Sack  "  Handb.,"  v.  Mracek,  p.  290, 

•^  Rosenbach,  Therap.  Alonafsh.,  1903. 

^  Leipzig,  med.  Monatsh.^  1907,  vi. 

''  Chemist  and  Druggist,  August  25,  1906. 


yi^  THE  ANIMAL   PARASITES   OF   MAN 


Pulex  irritans  (Human  Flea). 

The  bite  of  the  flea  produces  a  shght  discharge  of  blood  about  the 
size  of  a  pin's  head,  which  rapidly  becomes  surrounded  with  a  circular 
area  similar  to  a  patch  of  roseola.  The  redness  fades  away  after 
a  longer  or  shorter  while  (several  hours),  whilst  the  discharge  of  blood 
is  to  be  seen  for  one  or  two  days  longer.  In  dirty  people  the  whole 
body  mav  be  covered  with  such  discharges  of  blood.  Individuals 
with  very  delicate,  sensitive  skin,  especially  small  children,  show  true 
wheal  formation  at  the  site  of  the  bite.  In  certain  cases  there 
develops  from  one  such  single  bite  an  urticaria  that  extends  over 
a  large  part  of  the  body.  The  manner  by  which  an  irritating  sub- 
stance is  introduced  into  the  skin  upon  biting  by  the  bed  bug  and 
also  by  the  flea  is  clear.  The  bite  is  followed  by  a  feeling  of  itching, 
which  is  liable  to  rob  nervous  persons  of  their  sleep.  Sensitive 
individuals  are  upset  even  by  the  fleas  moving  over  the  surface  of  the 
skin  during  their  rest  at  night. 

Treatment  consists  in  extreme  cleanliness,  capture  of  the  parasites^ 
sprinkling  the  body  and  bed-linen  with  insect  powders.  The  fleas 
are  difficult  to  remove  from  barracks,  schools  and   hospitals. 


Dermatophilus  (Sarcopsylla)  penetrans  (Sand  Flea). 

The  fertilized  females  penetrate  into  the  skin  with  their  heads,  and 
here  they  swell,  in  consequence  of  the  numerous  and  growing  eggs 
and  larvae,  to  a  white  ball  the  size  of  a  small  pea,  on  which  the  head 
is  recognizable  only  as  a  small  brown  point. 

In  this  way  a  small  brown  tumour  arises,  over  which,  at  the 
commencement,  the  skin  is  not  reddened  ;  after  some  days,  however, 
it  becomes  inflamed;  in  the  centre  of  it  a  small  opening  is  seen. 
If  the  parasite  is  not  extracted  the  skin  that  lies  over  it  becomes 
destroyed  by  suppuration,  and  thus  becomes  removed.  At  the 
commencement  the  part  affected  itches,  with  increasing  inflammation  ; 
the  symptoms  of  irritation  become  more  severe  and  may  amount 
to  actual  pain.  If  the  small  suppurative  processes  be  neglected, 
inflammation  and  gangrenous  and  septic  processes  may  arise.  The 
region  of  the  body  sought  out  by  preference  by  the  sand  flea  is 
the  sole  of  the  foot,  the  toes,  under  the  free  ends  of  the  nails  and 
the  digito-plantoid  folds — more  rarely  the  scrotum,  thigh  and  other 
parts  are  attacked  (Scheube^).  The  number  of  parasites  found  on 
one  person  may  amount  to  several  hundreds. 


Scheube,  "Die  Krankh.  d.  warmer  Lander,"  1896. 


SUPPLEMENT  715 

Treatment  consists  in  the  removal  of  the  parasites  from  the  skin 
with  a  needle  or  a  small  sharp  knife  and  the  application  of  a  bandage. 
Rubbing  the  feet  with  copaiba  or  Peru  balsam,  sprinkling  them  with 
insect  powder,  or  washing  them  with  bay  rum  (Berger^)  acts  as  a 
prophylactic  or  removes  the  irritation  of  the  skin  produced  by  the 
parasites. 

Myiasis. 

Under  the  name  of  myiasis  we  designate  the  complex  symptoms 
which  parasitic  dipterous  larvae  give  rise  to  in  man  (Braun),  and  we 
conceive  under  the  term  myiasis  externa  (dermatosa  s.  cutanea)  all 
lesions  of  the  human  integument  caused  by  fly  larvae  and  of  the 
cavities  covered  with  mucosa  therewith  connected,  such  as  the 
external  auditory  meatus,  the  oro-nasal  cavity,  the  urethra  and  vagina. 
The  occurrence  of  dipterous  larvae  in  the  digestive  tract  is  named 
myiasis  intestinalia  or  interna. 

Myiasis  externa. 

The  larvae  of  a  species  of  fly  belonging  to  the  Muscidce,  Liicilia 
inacellaria,^  are  found  in  relative  frequency  in  the  nose,  especially 
in  America  and  India.^  Riley*  has  stated  that  the  screw-worm  of 
Central  America  and  of  the  United  States  is  nothing  else  than  the 
larva  of  Liicilia  inacellaria,  and  also  that  the  Brazilian  fly  named 
''  berna  "  may  be  no  other  than  Lncilia  macellaria.  Their  offspring 
may  set  up  inflammatory  disturbances  in  the  soft  tissues  of  man.  This 
fly  has  a  wide  distribution,  from  the  Argentine  Republic  to  Canada, 
also  in  the  British  portions  of  the  East  Indies,  where  the  disease  is 
named  "  peenash."  This  word  is  derived  from  the  Sanskrit,  and  is 
said  to  be  a  collective  name  for  all  diseases  of  the  nose.  Lahory^ 
states  that  w^ithin  a  period  of  nine  years  ninety-one  cases  of  " peenash '^ 
occurred  in  Allyghar,  two  of  these  ending  fatally.  Liicilia  macellaria 
is  not  at  all  timid  but  bold,  like  the  house-flies  and  blue-bottles, 
its  relatives.  It  not  only  lives  at  no  great  distance  from  human 
dwellings,  and  forces  its  way  into  villas  and  country  houses,  but  even 
attacks  its  victims  without  awaking  them  from  their  sleep.  Although 
this  species  shows  a  certain  preference  for  nasal  cavities  affected  with 
catarrh  or  pus  (v.  Frantzius^),  and  also  the  external  auditory  meatus, 

'  Berger,  Therap.  Monalsh.,  April,  1907. 

-    {Chrysomyia  macellaria,  p.  587. — F.  V.  T.] 

^  [C.  macellaria,  Fabricius,  the  screw- worm  fly,  is  found  in  tropical  America  and  the 
West  Indies.  The  genus  is  restricted  to  America.  The  species  from  India  is  a  Pycnosoma. — 
F.  V.  T.] 

^  Riley,  American  Naturalist,  1883,  xvii. 

•^  Lahory,  Edin.  Med.  lourn.,  1856. 

6  V.  Frantzius,   Virchow's  Archiv,  1868,  xliii. 

45 


7l6  THE   ANIMAL   PARASITES   OF   MAN 

as  well  as  ulcerated  or  wounded  parts  of  the  body,  and  even  badly 
ulcerated  skin  carcinoma  (Lutz^),  it  is  not  a  rare  thing  for  it  to 
penetrate  into  one  of  the  above-mentioned  cavities  rapidly  to  deposit 
its  eggs,  without  these  parts  having  been  previously  affected.  The 
report  also  of  Conil,*^  in  which  these  flics  bear  the  name  of  Calliphora 
anthropophaga,^  is  an  interesting  one.  Probably  it  was  the  same  species 
of  Muscid  in  the  cases  of  myiasis  nasi  observed  by  von  Tengemann, 
Delasiauve,*  Weber,^  Mankiewicz,^  and  Kirschmann.^  In  the  case 
recorded  by  Prima,^  and  in  that  recorded  by  Britton,^  the  issue  was 
a  fatal  one ;  in  the  latter  the  larvae  escaped  through  the  pharynx 
and  nose  ;  the  hyoid  bone  and  the  soft  parts  of  the  palate  were 
destroyed,  the  speech  and  power  of  swallowing  'were  hindered.  At 
the  post-mortetn  extensive  destruction  of  the  internal  nose  was  found, 
so  that  the  nasal  bones  could  only  be  kept  in  their  position  by  the 
aid  of  the  external  skin.  Even  during  life  227  larvae  escaped. 
Similar  destructive  processes  were  found  in  the  case  communicated 
by  Richardson. ^^  In  two  cases  reported  by  Schmidt^^  300  and  350 
larvae  were  respectively  removed  from  the  nose,  and  the  patients 
recovered.  Wolinz^^  found  his  patient  had  lost  consciousness,  and 
that  in  the  pus  filling  up  the  entrances  to  the  nose  numerous  larvae 
were  moving;  recovery  followed.  In  the  case  communicated  by 
Adler,^^  more  than  150  larvae  escaped  from  the  nose  of  an  old  man. 
Curran^*  states  that  people  suffering  from  ^'  peenash  "  frequently  die 
from  meningitis.  The  cases  reported  by  Pierre'^  related  to  the  forms 
of  severe  myiasis  frequently  to  be  observed  in  Guiana.  In  a  patient 
who  was  suffering  from  typhus  (  ?  typhoid),  Douglas^^  found  the 
conjunctival  sacs  full  of  larvae ;  in  two  other  individuals  the  nasal 
cavities  were  attacked. 

The  case  observed  by  Summa^^  was  that  of  a  man,  aged  28,  who 
suffered  from  nasal  obstruction,  foetor,  epistaxis  and  pain  in  the  nose. 
Out  of  seven  of  the  cases  occurring  at  Fort  Clark,  U.S.A.,  and  in  its 

'  Lu*z,  see  Joseph,  Deutsch.  med.  Zeitg.,  1885. 

*  Conil,  Annal.  de  Science  nat.  zooL,  1878. 

•  [This  fly  belongs  to  the  genus  Cordylobia,  and  is  peculiar  to  Africa,  C.  anthropophaga^ 
or  the  tumbri  fly,  is,  when  a  larva,  a  subcutaneous  parasite  of  man  and  animals. — F.  V.  T.] 

^  Delasiauve,  Gerhardt's  "Handb.  d.  Kinderkrankh.,"  1878,  iii. 
2  Weber,  Mexique  Rec.  d.  Mhn.  de  Med.  milit.^  1867. 
^  Mankiewicz,  Virchow's  Archtv,  1868,  xliv. 

'  Kirschmann,  Wien.  med.  Wochenschr.,  1881.  ^  Prima,   **  These  de  Paris,"  1881. 

^  Britton,  Cambridge,  Massachusetts,  1883. 

'"  Richardson,  Medical  Monthly,  1883.  "  Schmidt,  Texas  Med.  Journ.,  1887. 

'•-  Wolinz,  Wratsch,  1884.  i3  Adler,  Med.  Record,  1885. 

'*  Curran,  Med.  Press  and  Circ,  1887. 
'=*  Pierre,   "These  de  Paris,"  1888. 
'^  Douglas,  Kansas  City  Med.  Index,  1890. 
"  Summa,  St.  Louis,  1889. 


SUPPLEMENT  717 

neighbourhood,  six  ended  fatally  ;  in  all  these  cases  KimbalP  dia- 
gnosed ozaena  ;  attracted  by  the  strong  odour  the  flies  forced  their 
way  into  the  noses  of  the  patients  when  asleep  and  there  deposited 
their  ova.  In  a  case  reported  by  Carriere^  an  abscess  of  the  nasal 
septum  was  produced  by  the  larvae  of  flies  ;  Chiodi^  reports  seven 
cases  of  myiasis  due  to  Lucilia  macellaria  ;  among  these  was  a  case 
of  rhinitis  myiatica,  in  which  a  cerebral  abscess  leading  to  a  fatal 
termination  developed,  being  produced  by  the  migration  of  a  larva 
into  the  brain.  Among  the  three  cases  of  Lesbini^  was  that  of  a  girl, 
aged  16,  with  250  larvae  in  the  diseased  nasal  cavity.  Quintano^ 
observed  larvae  beneath  the  eyelids  in  one  case.  It  is  possible  that 
the  cases  of  Cesare^  and  Calamida''  w^ere  those  of  myiasis  nasi  due  to 
Lucilia  macellaria.  The  larvae  are  also  found  in  the  nasal  accessory 
sinuses,  as  is  seen  from  the  cases  reported  by  De  Saulle^  (frontal 
sinus),  Delasiauve^  (frontal  sinus),  MacGregor^^  (antrum  of  Highmore), 
and  Bordenave^^  (antrum  of  Highmore). 

If  a  survey  is  made  of  the  literature  of  the  cases  described  of 
myiasis  nasi  produced  by  Lucilia  macellaria^^  the  following  information 
is  forthcoming  :  In  Europe  this  form  of  the  disease  is  of  very  rare 
occurrence,  whilst  in  America  and  India^^  it  is  frequent.  Persons 
suffering  from  ozaena  are  rendered  the  most  liable  to  danger  as  the 
penetrating  odour  entices  the  flies  in  tropical  countries  with  intense 
frequency,  so  much  so  that  v.  Frantzius  does  not  consider  this 
myiasis  as  an  independent  disease,  but  as  a  complication  of  ozaena  of 
frequent  occurrence  in  warm  countries.  The  infection  is  so  far 
of  interest  in  its  nature,  in  that  it  only  takes  place  during  the  day. 
The  fly  is  on  the  wing  only  by  day  when  the  sun  is  shining,  and 
consequently  only  deposits  its  eggs  at  this  time.  Therefore  persons 
suffering  from  ozaena  are  principally  exposed  to  the  danger  of  being 
pursued  by  the  flies  when  they  succumb  to  sleep  during  the  mid-day 
hours  in  the  open  or  in  dwellings  that  are  not  closed  up. 

Headache  is  the  symptom  which  most  troubles  the  patients. 
It  extends  over  the  whole  cranium  and  persists  uninterruptedly,  with 


'  Kimball,  New  York  Med.  Journ.,  1893. 

2  Carriere,  Gaz.  hebd.  de  Med.  et  de  Chir.,  1898,  xciv. 

3  Chiodi,  La  Argent.  Med.,  March  i,  1905.  ^  Lesbini,  ibid. 
•^  Quintano,  *'  Cronic  oftalm.  de  Cadiz,"  1878. 

6  Cesare,  Arch.  ital.  di  Otol.,  April,  1903. 

■^  Calamida,  Giorn.  d.  A\  Accad.  de  Med.  di  Torino,  September,  1903. 

«  De  Saulh,  Gaz.  des  Hdp.,  Paris,  1857. 

»  Delasiauve,  Gaz.  hebd.  de  Med.,  Paris,  1885. 
^0  MacGregor,  Arch.  gin.  de  Med.,  No.  1,031. 

"  Bordenave,   **  Deuxieme  Mem.  presente  a  I'Acad.  de  Chir.,"  v,  p.  387. 
"   [And  the  other  species,  of  course,  must  be  included  here. — F.  V.  T.] 
'^   [Concerning  Europe  and  India,  macellaria  does  not  occur. — F.  V.  T.] 


7l8  THE   ANIMAL   PARASITES   OF   MAN 

more  or  less  severe  periods.  Violent  headaches  in  the  frontal  and 
buccal  regions  are  almost  always  present  in  this  complaint;  they  are 
experienced  either  only  on  one  side  or  on  both  simultaneously  ; 
sometimes  the  pain  is  extended  to  the  lower  jaw  and  region  of  the 
neck,  following  the  whole  extent  of  the  trigeminal  nerve.  The 
inflammation  of  the  nasal  mucosa  produced  by  the  penetration  into 
it  of  the  larvae  extends  right  into  the  frontal  sinus  and  antrum. 
Simultaneously  the  patients,  at  the  height  of  their  trouble,  suffer 
from  persistent  sleeplessness  and  severe  vertigo,  so  that  they  reel  and 
cannot  walk  straight ;  excessive  sneezing  always  sets  in  at  the  com- 
mencement. The  larvae  immediately  spread  over  the  nasal  mucosa  to 
seek  a  place  suitable  to  feed,  and  irritate  the  nasal  mucous  membrane 
by  the  tickling  sensation  they  produce.  Later  the  patients  frequently 
sneeze  when  the  maggots  move  to  and  fro. 

One  very  characteristic  symptom  consists  in  the  peculiar  swelling 
of  the  face,  which  is  extended  either  over  the  whole  or  only  one  half 
of  it,  and  may  alternate  with  attacks  of  erysipelas  (Brokaw^). 

The  discharge  from  the  nose  is  of  special  diagnostic  value.  It 
consists  of  a  blood-stained  serous  matter  or  blood-stained  fluid, 
which  is  perpetually  trickling  from  one  or  both  nostrils.  The  larvae 
especially  choose  the  anterior  portions  of  the  nasal  cavity,  where  they 
can  be  seen  lying  in  groups  together  at  the  base  of  the  choanae. 
The  consequence  of  this  is  that  the  soft  palate  becomes  intensely 
swollen,  and  this  in  turn  makes  swallowang  very  difficult  ;  speech  is 
impeded,  and  the  voice  acquires  a  nasal  intonation.  Symptoms  of 
fever  become  more  or  less  pronounced  according  to  the  number 
of  larvae  present,  and  according  to  the  nature  and  constitution  of  the 
individual.  The  appetite  is  in  abeyance  throughout  the  whole  dura- 
tion of  the  illness,  and  sometimes  there  is  the  onset  of  slight  attacks 
of   diarrhoea. 

If  the  larvae  are  not  removed  in  good  time  there  follows  excessive 
destruction  of  the  interior  of  the  nose  and  of  the  turbinals ;  and  the 
whole  nasal  framework  undergoes  disintegration,  frequently,  too,  the 
velum  palati,  so  that  the  larvae  come  into  sight  in  the  oral  cavity. 
Individuals  thus  severely  attacked  succumb  through  exhaustion, 
symptoms  of  meningitis  (cerebral  abscess)  or  septicaemia  (Prima"). 
Twenty-one  out  of  thirty-eight  cases  recorded  (collected)  by  Maillard^ 
died. 

The  method  of  prophylaxis  is  self-evident  from  what  has  been 
stated.     On  bright  summer  days  neither  the  healthy  nor  those  suffering 

'  Brokaw,  see  Seifert,  in  Heymann's  "  Handb.,"  p.  595. 

2  Prima,  "These  de  Paris,"  1881. 

^  Maillard,  "  These  de  Montpellier,"  1870. 


SUPPLEMENT  719 

from  diseases  of  the  nose  should  sleep  during  the  day-time  in  the 
open  or  in  public  habitations  :  sufferers  from  nasal  diseases  should 
pay  special  attention  to  this. 

Treatment  consists  in  the  removal  of  the  larvae  ;  this,  however,  is 
not  always  easy. 

With  regard  to  the  methods  which  have  proved  to  be  effectual  in 
the  destruction  of  living  larvae  and  their  expulsion  from  the  nose, 
strongly  smelling  and  easily  diluted  fluids  come  first,  such  as  alcohol, 
eau-de-Cologne,  and  ether,  which  should  kill  the  creatures  when 
injected  into  the  nostrils.  The  earlier  physicians,  such  as  Salzmann,^ 
Honold,^  and  Henkel,^  have  seen  good  results  from  the  use  of  these 
methods,  whilst  Mankiewicz''  and  Goldstein^  obtained  no  results 
whatever.  Kimball's^  careful  investigations  have  shown  that  a  decoc- 
tion of  bitter  herbs  recommended  by  Behrends''  (tansy,  wormwood) 
have  just  as  little  effect  as  the  tobacco  decoction  employed  by 
Boerhave^  and  Kilgour.-*  The  sternutatories  employed  by  the  older 
physicians  are  entirely  neglected.  Delasiauve'^  experienced  good 
results  from  the  inhalation  of  the  smoke  of  paper  cigarettes,  which 
were  soaked  with  a  solution  of  2-0  pot.  arsenic  in  30*0  distilled  water. 
Whilst,  according  to  Kimball,  balsam  of  Peru  had  no  effect  on  the 
larvae,  Mankiewicz  succeeded  in  removing  the  larvae  from  the  nose 
with  the  help  of  that  drug.  Turpentine  steam  or  mixtures  of  turpentine 
employed  by  Indian  physicians  have  not  been  very  effectual  according 
to  Moore,^^  Kimball  and  Goldstein.  Success  has  been  attained  in 
some  cases  by  the  use  of  insufflations  of  calomel  (Roura,^^  Cerna,^^ 
Schmidt^"*)  or  of  iodoform  (PascaP^).  Joseph^^  recommends  con- 
centrated alum  solution  being  sniffed  up  into  the  nose  as  very  effectual. 
Sublimate  and  carbol  solutions  do  not  appear  to  be  very  successful 
(Kimball,  Moore,  Goldstein),  whilst  benzine  inhalations  (Pierre^^) 
have  shown  better  results.  Scheppegrell^^  strongly  recommends 
injections  of  oil  which  kifl  the  larvae,  while  it  is  perfectly  harmless 
to  the  nasal  mucosa.  Cesare^^  employed  nasal  lavages  with  solutions 
of  salicylate  of  soda  with  good  results,  and  Calamida^^  lavages  with 
physiological  saline  solution.     Bresgen^^  recommends  the  nose  being 

'  Salzmann,  see  Tiedemann,  Mannheim,  1844.  "^  Honold,  ibid. 

3  Henkel,  ibid.  ^  Mankiewicz,  Virchov/s  Archiv,  1868,  xliv. 

*  Goldstein,  New  York  Med.  yourn.y  1892. 

6  Kimball,  ibid.,  1893. 

'  Behrends,  see  Tiedemann.  ^  Boerhave,  ibid. 

^  Kilgour,  ibid.  ^^  Ddtasiauve,  loc.  cit. 

'1  Moore,  Chicago  Med.  Times,  1893.  ^^  Roura,  Gaz.  di  San.  inilit.,  1884. 

•3  Cerna,  New  York  Med.  J  own.,  1893.  '^  Schmidt,  Texas  Courier,  1884. 

'5  Pascal,  Arch.  d.  Med.  milit.,  1895.  '^  Joseph,  Deutsch.  med.  Zeitg.,  1885. 

'7  Pierre,  "These  de  Paris,"   1888.         '»  Scheppegrell,  New  York  Med.  Journ.,  1898. 
'^  Cesare,  loc.  cit.  ^o  Calamida,  loc.  cit. 

2'  Bresgen,  Eulenburg's  "  Real.  Encyclopadie,"  third  edition. 


720  THE   ANIMAL   PARASITES    OF   MAN 

cocainized  and  the  larvae  being  removed  with  a  pincette.  Roorda- 
Smit^  cocainized  the  nose,  then  insufflated  calomel  and  plugged  the 
nose  with  a  gauze  tampon  dusted  with  calomel.  After  two  hours 
fifty-six  larvae  crawled  out  along  the  plug.  Continuation  of  the 
treatment  resulted  in  a  complete  cure. 

Injections  of  chloroform  water  (Jourdran^)  or  chloroform  inhala- 
tions, or  injections  of  pure  chloroform  into  the  nose,  have  proved 
the  most  effectual  (Goldstein,^  Osborn,^  Jourdran,  Durham,^ 
Jennings,^  Kimball/  Mackenzie,^  Oatmann/^  Zarniko,^^  Antony,^' 
Folkes^^).  Camphorated  carbolic  solutions  are  very  well  spoken  of  : 
Grayson^^  states  that  these  kill  the  larvae  immediately.  Some  authors 
have  removed  the  larvae  with  forceps  (Goldstein^'^),  others  with 
pincettes ;  thus  Brokaw  extracted  200  fragments  with  the  forceps, 
Pascal  eighty  fragments  with  the  pincettes,  and  Wolinz^^  also  appears 
to  have  removed  the  larvae  with  forceps. 

Greater  operative  measures  than  these  do  not  appear  to  have  been 
undertaken  in  latter  days ;  yet  Morgagni^*^  states  that  the  army 
surgeon,  Caesar  Mogatus,  at  Bologna,  first  trephined  the  frontal  sinus 
and  then  extracted  a  ''  worm  "  from  it. 

Larvae  of  other  Muscidce  have  come  under  observation  much  more 
rarely  (ChevaP'  [larvae  of  Galleria  melloiiella^^],  Bond,^^  DumesniP^ 
[larvae  of  PiopJiila  casei]).  Species  of  the  genus  Scolopendra 
{Myriapoda),  which  all  shun  the  light  and  seek  their  food  during  the 
night — which  consists  of  animal  and  vegetable  substances — frequently 
make  their  way  into  the  nasal  cavities  of  people  when  asleep.  They 
are  found  not  only  in  the  nose,  but  in  the  accessory  cavities.  In  the 
chapter  on  the  '*  Parasites  of  the  Nose  "^^  we  have  collected  striking 

'  Roorda-Smit,  Deutsch.  vied.  Wochenschr.,  ico6. 

^  Jourdran,  Aich.  de  Med.  nav.,  1895. 

^  Goldstein,  Neiv  York  Med.  Journ.,  1892. 

^  Osborn,  Daniel's  Mid.  Journ.^  1891. 

5  Durham,  Chicago  Med.  Times,  1893. 

^  Jennings,  Kansas  City  Med.  Index,  1890. 

"^  Kimball,  New  York  Med.  Journ. ^  1893. 

^  Mackenzie,  "  Diseases  of  the  Nose  and  Throat." 

^  Oatmann,  Med.  Mirror,  February,  1894. 
^°  Zarniko,  "  Lehrb,  d.  Krankh.  d.  Nase." 

'  Antony,  Bull,  Soc.  vied,  des  Hop.  de  Paris,  1903. 
'2  Folkes,  New  York  Med.  Record,  1907. 
'3  Grayson,  St.  Louis  Med.  and  Surg,  yonrn.,   1891. 
1*  Goldstein,  N'ew  York  Med.  Journ.,  1892. 
'^  Wolinz,  Wralsch,  1884.  '^  Morgagni,  see  Tiedemnnn. 

'^  Cheval,  Journ.  de  Med.  et  de  Chir.,  1893, 
'^   [This  is  the  larva  of  a  moth.  — F.  V.  T.] 
'*  Bond,  ////.  Zeniralbl.  f.  Laryng.,  1896. 
2°  Dumesnil,  see  Friedreich,  "  Die  Krankh.  d.  Nase,"  1858. 
^'  Seifert,  j^^  Heymann's  "Handb." 


SUPPLEMENT  72 I 

instances,  but  we  have  omitted  to  mention  the  observation  made  by 
Bertrand^  (Scolopendra  in  sinus  maxillaris)  and  that  made  by  Berg- 
mann^  (Scolopendra  in  sinus  frontahs).  In  the  same  chapter  some 
remarks  are  made  as  to  the  occurrence  in  the  nose  of  earwigs,  cater- 
pillars, scorpions  and  termites,  as  well  as  of  animals  which  have  not 
been  identified. 

The  larvae  that  develop  in  the  auditory  meatus  penetrate  the 
membrana  tympani,  destroy  the  middle  ear  and  may  produce  menin- 
gitis and  intracranial  suppurations.  In  one  case  Vesescu^  extracted 
seven  living  larvae  from  the  ear  with  the  aid  of  a  thin  pair  of  pincettes. 
Kohler^  recommends  the  infusion  of  drops  of  ol.  terebinth,  to  destroy 
the  larvae,  Ouintano^  the  insufflation  of  the  following  powder  :  Oxid, 
hydrarg.  rubr.,  sulfur.,  aa  I'o  grm.,  pulv.  gi.  arab.  8*0  grm.  ;  Lesbini^ 
recommends  tincture  of  iodine.  In  the  case  reported  by  Henneberg'' 
the  larvae  were  those  of  Lucilia  ca'sar. 

Eye  affections  due  to  Lucilia  iiiacellaria  are  very  uncommon  ;  the 
literature  relating  to  the  lesions  of  the  eye  produced  by  the  larvae  of 
flies  has  been  collected  in  Kayser's^  work.  In  the  cases  under  the 
observation  of  Schultz-Zeyden'**  both  the  eyes  of  a  female  tramp  were 
destroyed,  and  quantities  of  larvae  were  also  found  in  the  nasal  fossae 
and  in  the  ears. 

The  Lucilia  is  found  relatively  seldom  on  the  cutaneous  surface. 
Henneberg's^°  case  was  that  of  a  neglected  girl,  aged  20,  in  whom 
countless  larvae  (L.  ccesar)  were  found  in  a  plica  polonica  ;  after  the 
plica  polonica  had  been  removed  the  scalp  was  found  to  be  covered 
with  a  large  quantity  of  ulcers  which  swarmed  with  larvae,  large  and 
small.  The  skin  of  the  trunk  was  also  much  macerated  and  covered 
w^ith  larvae.  Death  resulted  from  sepsis ;  Westenhoffer"  remarks  on 
this  case  that  a  lesion  of  the  head  from  which  the  patient  had  suffered 
previously  and  the  perpetual  state  of  intoxication  in  which  she  was 
had  probably  given  rise  to  the  lodgment  of  the  fly  larvae.  Whether 
the  communications  made  by  Munk^^  Qf  maggots  in  the  mouth 
relate  to  Lucilia  I  do  not  know.     Vesescu,^^  in  one  case  with  extensive 


'  Bertrand,  Soc.  med.  de  Bologne,  1839. 

2  Bergmann,  Koi-respondenzbl.  d.  deiilsch.  Ges.  f.  Psych.,  Neuwied,  1 859. 

*  Vesescu,  Riv.  sliintelor  vied.,  February,  1906. 

*  Kohler,  Monatsschr.  f.  Ohretiheilk.,  1885. 

*  Quintano,  see  Seifert,  loc.  cit. 

^  Lesbini,  La  Argent.  Med.,  1905. 

■^  Henneberg,  Berl.  med.  Ges,,  February  18,  1903. 

^  Kayser,  Kliu.  Monatsbl.  f.  Augenheilk.,  1905. 

^  Schultz-Zeyden,  ^^r/.  i/z'«.  Wochenschr.,  1906. 
^^  Henneberg,  Berl.  med.  Ges.,  February  18,  1903. 
^'  Westenhoffer,  Verein  f.  innere  Med.,  Berlin,  May  7,  1906. 
^2  Munk,  Wien.  med.  Presse,  xxi. 
*^  Vesescu,  loc.  cit. 


72  2  THE   ANIMAL   PARASITES   OF   MAN 

ulceration  and  deep  fistulae  in  the  skin,  removed  176  larv^e  with  the 
pincette.  In  Roorda-Smit's^  case  there  were  two  ulcers  in  the  neck 
of  a  girl,  aged  17,  and  larvae  appeared  at  their  base.  After  dusting 
with  calomel  and  the  appUcation  of  a  bandage  the  next  day  fifty-two 
dead  or  half-dead  larvae  came  to  light.  Recovery  took  place. 
Lesbini/  in  the  case  of  an  old  lady,  saw  numerous  larvae  in  an  ulcer 
of  the  leg  she  was  suffering  from.  Hector's^  case  appears  to  have 
been  one  of  myiasis  cutanea  provoked  by  Lucilia. 

The  first  exact  observations  of  myiasis  cutanea  from  SarcopJiaga 
magnifica  are  due  to  Wohlfahrt,*  in  whose  honour  Portschinsky'"' 
named  this  species  of  fly  S.  wohlfahrti.  Portschinsky  ascertained  that 
S.  ivohlfahrii  was  not  confined  to  man  as  its  sole  host,  but  that  several 
of  our  domestic  animals,  such  as  cattle,  horses,  pigs,  dogs  and  geese, 
w^ere  visited.  In  these  animals  small  wounds  serve  to  entice  the  flies 
and  to  supply  them  with  a  suitable  site  for  the  deposition  of  their 
eggs.  The  oral  armature  of  the  young  larvae  renders  it  easy  for  them 
to  penetrate  not  only  the  mucosa  and  cutaneous  surface  but  also 
intact  places  in  the  submucous  connective  tissue.  In  many  localities 
more  than  half  the  herds  have  proved  to  be  infected  by  the  flies. 
The  fly  only  frequents  open  spaces  and  never  enters  human  dwellings, 
and  is  so  timid  that  it  approaches  man  only  during  sleep  ;  infection, 
therefore,  takes  place  only  out  of  doors,  in  summer,  in  clear,  warm 
weather,  and  only  in  such  individuals  as  sleep  in  the  open  air. 
Individuals  are  most  exposed  to  risk  who  suffer  from  catarrhs  or 
inflammations,  combined  with  purulent  secretions  of  the  nasal  cavity 
(ozaena),  or  otorrhcea,  or  ulcers  in  any  parts  of  the  body  accessible  to 
the  female  fly. 

The  frequency  and  intensity  of  the  infection  will  be  in  inverse 
proportion  to  the  advance  in  civilization  of  the  inhabitants,  their  idea 
of  cleanliness,  their  having  timely  medical  aid  and  the  chances  of  their 
being  rapidly  attended  to.  On  that  account  the  majority  of  cases  of 
myiasis  (Sarcophaga)  are  reported  from  Russia.  The  literature  of  this 
kind  of  myiasis  nasalis  is  not  very  extensive  ;  in  addition  to  Wohlfahrt, 
Portschinsky  and  Joseph,*  there  is  a  communication  by  Gerstacker,^ 
who  found  fifteen  adult  larvae  of  5.  wohlfahrti  in  the  nasal  cavity  of 
one  man.  The  larvae  transmitted  from  Ordruf  by  Dr.  Thomas  to 
Low,^  in  Vienna,  which  were  discharged  from  the  nose  of  a  woman, 


'  Roorda-Smit,  Deutsche  med.  Wockenschr.^  1906. 

^  Lesbini,  loc.  cit. 

'  Hector,  Lancet^  1902. 

*  Wohlfahrt,   "  De  vermibus  per  nares  excretis,"  Norimbergae,  1770. 

*  Portschinsky,  "Norae  Soc.  entomolog.  Rossicae,"  1875. 

*  Joseph,  Deutsch.  vied.  Zeitg.,  1885. 

'  Gerstacker,  "Sitzungsberichte  d.  Ges.  f.  naturf.  Freunde  in  Berlin,"  1875. 
"  Low,  Wien.  med.  Wochenschr,,  1883,  xxxi. 


SUPPLEMENT  723 

aged  71,  suffering  from  czaena,  were  recognized  by  the  well-known 
dipterologist  Braun  as  belonging  to  S.  wohlfahrti.  Among  the  cases 
reported  by  Joseph,  one  only  affected  the  nose;  it  was  that  of  a 
peasant  girl,  aged  11,  who  had  suffered  from  ozaena;  she  had 
travelled  on  the  open  road  and  had  there  gone  to  sleep.  Severe 
symptoms  set  in  and  death  followed  under  delirium.  In  making  the 
post-mortem  it  was  found  that  the  interior  of  the  nose  was  extensively 
destroyed  by  larvae  of  S.  wohlfahrti.  Powell  found  Sarcophaga 
larvae  in  two  persons  who  had  slept  in  the  open  air  ;  the  larvae  were 
killed  by  injections  of  chloroform  and  sublimate.  Destruction  of  the 
eyes  by  S.  wohlfahrti  has  only  been  observed  in  a  few  cases  ;  it  is 
reported  by  Cloquet^  that,  in  the  case  of  a  ragman  who  had  lain 
some  time  in  the  fields,  both  eyes  were  pierced  by  larvae.  On  the  outer 
skin  the  larvae  of  S.  wohlfahrti  have  been  found  more  than  once  in 
inflammatory  or  festering  areas.  Freund^  demonstrated  that  from 
a  live  year  old  child,  which  had  suffered  for  some  time  from  an 
impetiginous  eczema  of  the  skin  of  the  head,  from  two  suppurating 
abscess  cavities  which  extended  to  the  periosteum,  which  was  already 
affected,  twenty-one  living  larvae  were  taken  ;  rapid  healing  took  place 
under  antiseptic  bandaging. 

The  small  treatise  by  Balzer  and  Schimpff^  contains  two  new 
observations  on  myiasis  externa  ;  in  the  one  case  an  ulcer  on  a  man's 
foot  was  full  of  larvae,  in  the  other  case  thehead  of  a  woman  showed 
numerous  larvae  without  the  skin  of  the  head  being  destroyed. 
Brandt's'*  observation  is  interesting,  for  he  found  such  larvae  in  the 
gums  of  a  sick  person. 

The  impression  which  one  obtains  of  the  active  movement  of 
larvae  on  wounds  is  a  strange  and  at  the  same  time  uncanny  one. 
One  finds  that  the  larvae  to  obtain  protection  against  the  drying  of 
the  surface  of  the  abscess  almost  incessantly  burrow  with  their  heads, 
first  contracting  and  then  expanding  the  body,  which  rises  and  falls, 
and  keeping  the  tail  upwards.  Owing  to  these  movements  producing 
irritation,  increase  of  inflammation  may  ultimately  arise,  causing 
erysipelas  and  cellulitis. 

The  treatment  of  myiasis  nasalis  caused  by  Sarcophaga  is  the  same 
as  in  myiasis  caused  by  Lucilia,  and  in  the  other  places  where  found  it  is 
merely  a  question  of  the  removal  of  the  larvae  and  the  subsequent 
proper  treatment  of  the  surface  of  the  abscess.     In  Northern  Nigeria 


*  Cloquet,  see  Schultz-Zehden,  loc.  cit. 

"^  Freund,  Ges.  f.  innere  Med.  in  Wien,  December  5,  1901  ;  and  Wien.  med.  Wochenschr . ^ 
1910,  li. 

3  Balzer  and  Schimpff,  Annal.  de  Derm,  et  de  Syph.y  1902. 
<  Brandt,  Wratsch,  1888. 


724  THE    ANIMAL   PARASITES   OF   MAN 

Lelean^  found  Anchmeromyia  depressa  to  be  the  cause  of  myiasis 
externa.^ 

The  occurrence  of  Oestiid  larv^  in  a  human  being  is  very  rare, 
at  least  up  till  now  myiasis  oestrosa  has  been  very  seldom  observed 
in  man  in  Europe.  Whilst  the  hosts  of  the  Muscidct  comprise  a 
considerable  number  of  warm-blooded  animals,  on  which  the  larvae 
develop,  each  species  of  the  Oestridce  appears,  on  the  other  hand,  to 
have  a  definite  host  or  some  definite  hosts  of  the  class  Mammalia. 
No  species  of  Oestrid  is  peculiar  to  man.  Although  in  America,  as 
well  as  in  Europe,  Oestrus  lioniiuis  was  spoken  of  up  to  the  middle  of 
the  last  century,  no  such  species  exists. 

But  in  both  hemispheres,  in  America  much  more  often  than  in 
Europe,  Oestrid  larvae  have  been  found  in  man.  In  Florida,  Mexico, 
New  Granada,  Argentina,  Brazil,  Costa  Rica  and  other  districts, 
and  especially  where  large  herds  of  cattle  are  kept,  myiasis  oestrosa 
has  been  observed  in  shepherds,  huntsmen  and  amongst  the  rural 
population.  The  larvae  of  Hypodenna  hovls,  according  to  the  obser- 
vations of  Goudot,^  occur  as  a  parasite  in  man.  Poilroux^  found  larvae 
of  cavicolous  Oestridce  in  the  nose  of  a  man,  aged  55.  Amongst 
the  species  of  warble  flies,  whose  larvae  are  parasites  in  domestic 
animals  and  game  in  Europe,  reliable  observers  have  found  larvae  of 
two  kinds,  Hypodenna  bovis  and  Hypoderma  diana,  also  in  man.^ 

The  larvae  of  H.  bovis  have  very  seldom  been  observed  in  the  nose. 
The .  case  quoted  by  Kirschmann,*^  which  was  that  of  a  peasant 
woman,  aged  50,  who  was  suffering  from  ozaena,  and  in  which 
violent  attacks  of  sneezing,  epistaxis,  pain  in  the  forehead,  and  swelling 
of  the  face  were  observed,  is,  according  to  Low^  and  Joseph,^  not 
an  Oestrid  ;  Muscid  larvae  were  evidently  the  cause.  By  the  injection 
of  diluted  iron  chloride  solution  seventy-nine  larvae  were  removed 
from  the  nose.  In  the  case  reported  by  Kazoux^  the  species 
of  larva  is  not  definitely  known — at  least,  v.  Frantzius^^  did  not 
consider  them  Oestrid  larvae.  Joseph  does  not  definitely  say  that 
Oestrid  larvae  were  the  cause  of  a  case  which  he  quotes.  He  was  sent 
a  number  of  uninjured  larvae  of  Oestrus  ovis  ready  to  pupate,  which 


'  Lelean,  Bril.  Med.  Journ.^  1904. 

2   [Numerous  instances  of  attacks  by  Auchmeromyia  are  known  and  referred  to  under  that 
genus,  pp.  593-4'     The  species  referred  to  here  is  not  depressa.  Walker. — F.  V.  T.] 
•''  Goudot,  Annul,  d.  Set.  itat.,  1845. 
^  Poilroux,  Joiirn  de  Mid.,  Chir.,  etc.,  1809. 

^   \Hypodertna  linearis  is  frequently  confused  with  H.  bovis. — F.  V.  T.] 
6  Kirschmann,  Wien.  med.  Wochenschr.,  1881. 

■^  Low,  Wien,  med.  Wochenschr.,  1882.  ^  Joseph,  Deiitsch.  vied,  Zeitg.^  1885. 

^  Razoux,  Journ.  de  Mid,  Chir.,  etc.,  1 758. 
*°  V.  Frantzius,   Virchow's  Archiv,  1868,  xliii. 


SUPPLEMENT  725 

were  said  to  have  been  expelled,  during  violent  sneezing,  from  the 
nose  of  a  peasant  woman  who  had  suffered  for  six  months  from 
continuous  frontal  headache  and  chronic  nasal  catarrh. 

The  Oestrides  prefer  to  use  the  surfaces  of  wounds  on  the  skin  of 
man  to  lay  their  eggs,  which  develop  into  larvae  ;  but  they  often  use 
their  ovipositors^  to  make  a  fresh  wound.  In  this  case  there  arise  in 
the  skin,  and  particularly  in  the  subcutaneous  connective  tissue  of  the 
neck,  in  the  region  of  the  shoulder,  as  wxU  as  in  other  parts  of  the 
body  painful,  furuncle-like  inflammations  which  are  known  under 
the  name  of  gad-fly  boils.  These  boils  may  become  the  size  of  pigeons' 
eggs  ;  if  several  are  together,  they  appear  to  form  a  connected  tumour. 
Each  tumour  is  elastic  and  somewhat  movable,  and  has  an  orifice 
through  which  the  larva  breathes  and  discharges  its  excreta.  At 
times  these  turn  to  festers  and  gangrenous  disintegrations,  which  may 
even  cause  the  loss  of  a  limb.  Wilms^  had  the  opportunity  a  few 
years  ago  of  observing  a  case  of  myiasis  dermatosa  oestrosa  in  Leipzig. 
The  fistula  which  led  to  the  larva  was  slit  open  and  the  larva  extracted. 
As  a  notable  characteristic  of  myiasis  oestrosa  Joseph  states  that  the 
larva3  grow  very  slowly.  The  flight  time  of  the  Oestridcv  is  the  hot 
summer  months. 

Adams^  observed  on  the  Isthmus  of  Panama  a  number  of  cases 
of  a  skin  disease  which  is  caused  by  the  larvae  of  Dermatobia  noxialis 
{Gtisano-peliido-Muche).  The  larvae  penetrate  not  only  the  skin  but 
also  the  mucous  membrane  of  the  pharynx  and  larynx,  and  from  there 
proceed  through  the  tissue  to  the  subcutaneous  cellular  tissue.  The 
infection  seems  to  result  from  bathing. 

The  study  of  ^'  thimni,"  a  human  myiasis  caused  by  Oestrus  ovis, 
by  Ed.  and  Et.  Sergent,^  deals  more  with  the  zoology  and  with  the 
geographical  distribution  of  this  insect  in  North  Africa  than  with  the 
clinical  appearances  of  myiasis.  [This  paper  deals  with  matters  of 
great  interest,  with  important  facts. — F.  V.  T.] 

The  treatment  consists  in  the  removal  of  the  larvae  (from  the 
nose) ;  in  Brazil  it  is  the  custom  to  drop  tobacco  juice  into  the 
boil  in  order  to  kill  the  larvae  (Strauch^). 

One  is  only  justified  in  speaking  of  myiasis  intestinalis  when  there 
is  no  doubt  that  living  fly  maggots  or  flies  themselves  can  be  proved 
to  have  been  found  in  the  fresh  contents  of  the  stomach  or  intestine 


^  [The  O.  strides  appear  to  lay  their  ova  on  the  hair  of  animals.     They  do  not  puncture 
the  skin.-F.  V.  T.] 

-  Wilms,  Deutsch.  med.   Wochensclu:,  1897. 

8  Adams,  [our7t.  Atner.  Med.  Asscc,  1904. 

^  Ed.  and  Et.  Sergent,  Annal.  de  VInst.  Pasteur,  1907. 

^  Strauch,y^//;v/.  of  Cut.  Dis.,  1906. 


726  THE   ANIMAL   PARASITES   OF   MAN 

(Schlesinger  and  Weichselbaum^).  In  the  discussion  of  myiasis 
intestinalis  we  give  the  evidence  of  Schlesinger  and  Weichselbaum, 
as  well  as  that  of  Wirsing,^  to  which  must  be  added  a  number 
of  other  investigations. 

In  a  great  number  of  acute  cases  apparently  only  the  stomach  was 
affected,  there  being  no  signs  in  the  intestine.  In  these  cases  sudden 
illness  is  noticed,  colic,  sometimes  unbearable  pains  in  the  region  of 
the  stomach,  pyrosis,  vomiting  or  continuous  intense  inclination  to 
vomit,  occasionally  even  with  the  mixture  of  blood.  Frequently  a 
general  feeling  of  malaise,  twinges  of  pain  in  the  muscles,  and  attacks 
of  giddiness  were  notified,  very  rarely  fever.  Generally  all  the 
symptoms  disappeared  in  a  short  lime  when  the  larvae  had  been 
removed  by  an  act  of  vomiting  or  by  washing  out  the  stomach. 

It  is  well  to  note  that  in  the  history  of  many  cases  the  pains 
preceding  the  expulsion  of  the  larvae  are  stated  to  be  extremely 
violent. 

Acute  myiasis  of  the  intestinal  canal  frequently  runs  a  course 
without  special  symptoms  and  is  only  an  accidental  condition  ;  one 
.  has,  however,  in  such  cases  to  guard  against  errors.  The  faeces  may 
be  deposited  in  vessels  or  places  where  fly  larvae  are  in  great  numbers, 
or  a  subsequent  infection  of  the  faeces  with  the  eggs  or  larvae  of  flies 
may  have  taken  place.  Only  when  the  inspection  of  the  excrement 
immediately  following  defaecation  proves  the  presence  of  living  larvae, 
and  when  there  were  certainly  no  fly  larvae  in  the  vessel  previously, 
can  one  speak  of  the  passing  of  fly  larvae  from  the  intestine.  More 
frequent  than  the  cases  showing  no  special  symptoms  are  those  with 
pronounced  disturbances  in  the  intestinal  passage,  obstruction  or 
diarrhoea  (also  constipation  and  diarrhoea  alternately),  violent  and 
sometimes  agonizing  abdominal  pains  (Pottiez^),  which  preceded 
the  evacuation  of  the  larvae  and  subsided  after  their  removal.  General 
symptoms,  like  weakness,  languor,  transitory  vague  pains,  loss  of 
appetite,  sickness,  rarely  fever,  giddiness,  attacks  of  faintness,  epileptic 
attacks  (Krause*)  are  observed.  In  a  few  cases  blood  and  pus  have 
been  noticed  in  the  evacuation  of  the  bowels. 

In  the  cases  of  chronic  myiasis  of  the  intestine  the  aspect  of  the 
disease  is  dominated  by  the  complex  symptom  of  colitis  mucosa. 

The  following  features  are  noticeable,  namely,  the  intermittent 
passing  of  blood,  the  influence  over  the  expulsion  of  the  larvae  of 
mechanical  procedure  (massaging  of  the  abdomen),  the  duration  of 
the  process  for  several   years,  the  sometimes  enormous   number  of 

1  Schlesinger  and  Weichselbaum,  Wien.  klin.  Wochenschr.,  1902,  i. 

2  Wirsing,  Zeituhr.f.  klin.  Med.,  1906,  Ix. 

3  Pottiez,  Bull,  de  V Acad,  royale  de  Med.  de  Belgique,  xv. 
^  Krause,  Detiisch.  med.  Wochenschr.,  1886,  xvii. 


SUPPLEMENT 


727 


insects  contained  in  the  dejecta.  Another  cHnically  important  factor 
is  the  passing  of  the  larvae  in  batches.  While  for  some  time  no  larvcC 
may  appear  in  the  stools,  they  may  suddenly  be  ejected  in  great 
numbers,  either  because  the  conditions  of  feeding  are  not  suitable, 
or  because  medicaments  remove  them  from  the  intestine.  The 
haemorrhage  is  ascribed  by  Schlesinger  and  Weichselbaum  directly 
to  lesions  of  the  mucous  membrane  caused  by  the  larvae  ;  in  the  case 
reported  by  these  writers  there  were  found  shreds  of  tissue  as  well  as 
pus  in  the  stool.  The  pains  occurring  spontaneously  in  the  abdomen 
are  at  times  influenced  by  position  and  attitude  of  the  body,  often 
they  were  more  violent  after  rest  and  after  evacuation  of  the  bowels  ; 
often  they  were  continuous,  but  in  that  case  less  intense  ;  pressure  on 
the  abdomen  is  generally  little  felt.  The  condition  of  the  blood  was 
in  two  cases  (Pasquale^  and  Schlesinger  and  Weichselbaum)  a  marked 
chlorotic  one.  The  state  of  nutrition  seems  almost  always  to  suffer 
with  prolongation  of  the  disease,  but  in  Peiper's^  cases  this  was  not 
so.  The  condition  of  the  appetite  was  in  some  instances  good,  in 
others  very  bad.  A  frequent  symptom  is  headache  of  a  migraine-like 
character  and  neuralgic  pains  in  different  parts. 

Schlesinger  and  Weichselbaum's  case  shows  that  there  are  forms 
of  myiasis  intestinalis  which,  after  prolonged  sickness,  lead  to  death, 
and  that  in  consequence  of  the  formation  of  intestinal  abscesses 
stricture  of  the  intestine  may  arise  from  the  subsequent  formation  of 
a  scar. 

The  question  of  the  mode  of  infection  is  interesting ;  in  this 
mouth,  nose  and  anus  must  be  considered.  The  most  frequent  way 
is  certainly  by  means  of  food  on  which  flies  have  laid  their  eggs,  or 
which  is  permeated  with  young  maggots.  This  may  be  raw  (especially 
grated)  meat,  cheese,  fruit,  salad,  milk,  cabbage,  cold  farinaceous 
foods,  raspberries.  When  the  stomach  is  affected,  when  the  gastric 
juice  has  lost  acidity  and  power  of  digestion,  the  larvae  will  be  able 
to  stay  and  develop  more  easily.  According  to  Csokor,^  if  the  eggs 
get  into  the  gastro-intestinal  canal  of  man  with  the  food,  the  delicate 
stages  of  the  young  larvae  would  certainly  not  surviv^e  the  action  of 
the  gastric  juice.  Salzmann^  assumed  that  the  invasion  occasionally 
occurred  through  the  rectum,  the  larvae  creeping  into  the  anus  while 
the  person  is  asleep.  Wirsing  accepts  this  method  of  infection  for 
two  of  his  cases,  where  it  was  a  question  of  the  infection  of  an  infant. 
Salzmann*  reports  a  case  where  the  maggots  of  Anthomyia^  scalaris 

1  Pasquale,  Centralbl.f.  Bakt.,  1891. 

2  Peiper,  "  Fliegenlarv.  als  gelegentl.  Parasiten  d.  Menschen,"  Berlin,  1900. 

3  Csokor,  Wieti.  klin.  Wochenschr.,  1901,  p.   129. 

4  Salzmann,  Wiirttemberg.  med.  Korrespondenzbl.  1883,  liii. 

3   [This  is  presumably  Homalomyia  [FaTinia)  scalaris, — F.  V.  T.] 


728  THE    ANIMAL    PARASITES    OF    MAN 

were  passed  in  great  numbers  from  the  urethra  of  an  old  man.  The 
patient  had  been  catheterized  on  account  of  urethral  stricture  and  was 
probably  infected  with  eggs  or  larvae  at  the  same  time. 

The  diagnosis  of  the  affection  is  easy  and  sure,  if  Hving  larva'  are 
found  in  the  contents  of  the  stomach  or  in  the  stools,  and  if  contami- 
nation is  out  of  the  question. 

The  number  of  different  species  of  flies  whose  larva3  are  found  in 
myiasis  intestinalis  is  considerable.  The  larvae  of  species  of  Anthomyia 
(.4.  canicularis,^  A.  scalaris,  etc.),  of  Sarcophaga  carnaria  and 
6\  niagnifica  and  of  Mnsca  vomitoria^  are  especially  observed. 

The  prognosis  is  certainly  generally  favourable,  but  must  be  made 
with  some  reserve  in  chronic  cases,  in  view  of  the  observations  of 
Schlesinger  and  Weichselbaum  (intestinal  stenosis). 

The  treatment  must  aim  at  removing  the  larvae  as  soon  as  possible 
from  the  digestive  canal. 

In  cases  of  myiasis  of  the  stomach,  a  thorough  washing  out  of  the 
stomach  (Joseph,^  Staniek*)  is  to  be  preferred  to  emetics  used  with 
success  in  individual  instances ;  perhaps  it  would  be  advisable  to  add 
menthol  or  thymol  to  the  mixture. 

In  myiasis  of  the  intestine  internal  remedies  and  local  treatment 
of  the  intestine  must  be  considered. 

So  far  santonin  seems  to  have  proved  to  be  the  best  remedy. 
In  some  cases  extract,  filicis  maris,  calomel,  semina  cucurbitae, 
naphthalene  o'l  to  0*5  (Peiper^),  infus.  of  Persian  insect  powder 
(5  in  200),  mineral  waters,  Carlsbad  water,  seem  to  have  had  good 
results. 

P'or  irrigation  of  the  rectum,  weak  solutions  of  argentum  nitricum, 
tannin,  thymol,  gelatine,  ol.  ricini,  naphthalene  may  be  used.  Wirsing 
administered  an  aperient  (Rurella  compound  liquorice  powder)  and 
a  soap  enema  after  the  passing  of  the  first  larvae. 

The  principal  thing  is  the  prophylaxis,  which  must  include  the 
careful  protection  of  articles  of  food,  on  which  flies  may  lay  their 
eggs  (protection  by  glass  dishes,  tulle  or  fine  wire  nets).  Fruit 
should  not  be  eaten  before  being  washed  or  rubbed  with  a  cloth. 


^  [This  fly,  common  in  houses,  is  known  as  Homalomyia  canicularis,  and  the  next  belongs 
to  the  same  genus. — F.  V.  T.] 

2  [This  fly  belongs  to  the  genus  Calliphora,  not  Musca.—Y.  V.  T.] 

5  Joseph,  Deiitsch.  vied.  Zei(g.,  1885  and  1887. 

^  Staniek,  see  Schlesinger  and  Weichselbaum,  p.  47. 

«  Peiper,  "  Fliegenlarv.  als  gelegentl.  Parasiten  d.  Menschen,"  Berlin,  1900. 


SUPPLEMENT  729 

Gastricolous  Oestridae  (Creeping  Disease). 

Syn.  :  Creeping  eruptio7i;  Larva  migrans  ;  H autmaulwurf ;  Dermatomyiasis 
linearis  migrans  oestrosa ;  Hy ponomoderma  ;  Dermatitis  linearis  migrans ;  Linea 
migrans;  Epidermiditis  linearis  migrans  Wolossatik j  Kriechkrankheit ;  Haut- 
kratzschorf  ;  Myiase  hypodermique. 

Under  the  name  ^'  creeping  disease,"  R.  J.  Lee^  has  recorded 
a  peculiar  affection  of  the  skin  in  a  three  year  old  girl,  which  appeared 
first  in  the  form  of  pale  red,  thread-like  irregular  protuberances, 
which  seemed  partly  to  become  entwined  on  the  right  malleolus 
and  had  spread  without  causing  special  disturbances  to  the  abdomen. 
Dickinson,  Fox  and  Duckworth^  reported,  in  connection  with  this, 
that  they  observed  a  growth  of  this  red  line  of  about  i  in.  per 
diem.  Since  then  a  number  of  similar  cases  have  been  reported 
which,  without  doubt,  were  cases  of  larvae  creeping  under  the 
skin.  Crocker^  saw  such  a  case  in  a  two  year  old  girl,  the  progress 
of  the  red  line  varying  in  one  night  between  4  and  7J  in.  In  Europe 
the  first  case  was  observed  in  Vienna,  by  v.  Neumann  and  Rille,"^ 
also  in  a  two  year  old  girl. 

V.  Samson-Himmelstjerna,^  Sokoloff,*^  Rawnitzky'  found  larvae 
at  the  end  of  the  tract,  which  had  been  recorded  as  larvae  of  Gastro- 
philus  by  Cholodowsky.^  According  to  Blanchard  {Arch.  f.  Par., 
190 1 )  the  larvae  were  those   of  Hypoderma  hovis. 

How  these  larvae  get  into  the  skin  has  not  yet  been  definitely 
ascertained  ;  v.  Samson  is  of  the  opinion  that  they  usually  obtain 
access  to  man  as  larvae,  Stelwagon^  believes  that  the  infection 
generally  occurs  in  a  seaside  watering  place;  a  patient  of 
EhrmannV^  fell  ill  when  he  returned  from  the  manoeuvres,  where 
he  had  lain  for  some  time  on  the  ground.  Here  and  there  it  is 
reported  that  the  eruption  was  preceded  for  a  longer  or  shorter  time 
by  lesions  of  the  skin  (incised  wounds,  furuncles,  slight  excoriations, 
V.  Harlingen"). 

Twice  it  has  been  suggested  that  perhaps  the  parasites  might  come 
from  vineyard  snails  (Crocker,    Lenglet   and  Delaunay'^),    and  it  is 


»  R.  J.  Lee,/<7Mm.  Clin.  Soc:  Lond.,  November  27,  1874. 

2  Dickinson,  Fox  and  Duckworth,  ibid.^  1875. 

3  Crocker,  "  Diseases  of  the  Skin,"  1893  ;  **  Atlas  of  the  Diseases  of  the  Skin." 

*  V.  Neumann  and  Rille,  Wten.  klin.  Wochenschr. ,  1895  ;  Dermatologenkongr.,  Graz,  1895. 

*  V.  Samson-Himmelstjerna,  Wratsch,  1895;  Arch.f.  Derm.  u.  Syph.,  i897- 
«  Sokoloff,  Wratsch,  1896. 

'^  Rawnitzky,  Derm.  Zeitschr.,  v,  p.  704. 

«  Cholodowsky,  Wratsch,  1896. 

'  Stelwagon,  Journ.  Cut.  Dis.,  xxii,  8. 
'»  Ehrmann,  Wien.  derm.  Ges.,  November  17,  1897. 
"  V.  Harlingen,  Amer.  Journ.  of  Med.  Set.,  1902. 
12  Lenglet  and  Delaunay,  Annal.  de  Derm,  et  de  Syph.,  1904. 


730 


THE   ANIMAL   PARASITEvS   OF   MAN 


pointed  out  by  v.  Samson  that  in  Russia  the  infection  of  peasants 
who  work  in  the  fields  was  specially  frequent.  It  is  noticeable  how 
frequently  the  affection  begins  on  uncovered  parts  of  the  body  (face, 
hands,  arms)  ;  but  that  fact,  on  the  whole,  is  not  in  conflict  with  the 
statement  (Kengsep')  that  the  disease  makes  its  first  appearance 
over  the  nates,  because  children  often  sit  on  the  ground  and  play 
with  that  part  of  their  body  uncovered.  A  case  observed  by  us  was 
that  of  an  elderly  lady  who  did  not  do  this  and  was  properly  clothed, 
yet  showed  the  typical  lines  of  creeping  disease  on  the  nates,  and 
asserted  again  and  again  that  she  had  the  feeling  as  if  a  worm  were 
creeping  under  her  skin. 

The  disease  occurs  in  children  as  well  as  adults,  so  that  age,  sex 
and  calling  offer  no  determining  point  etiologically. 

The  clinical  symptoms  of  the  disease  consist  in  the  sudden 
appearance  of  itching  and  burning  ;  if  the  cause  is  looked  for  one 
perceives  a  red  line,  raised  but  little  above  th^^  surface  of  the  skin, 
with  irregular  curves,  never  branched,  but  often  entwined,  broadening 
more  or  less  rapidly  at  one  end  (i  to  15  cm.  in  twenty-four  hours). 
The  larva  can  be  seen  sometimes  with  a  lens  under  pressure  of  the 
skin  as  a  dark  spot ;  formations  of  pus,  such  as  other  larvae  produce, 
are  not  noticed ;  now  and  again  there  is  a  formation  of  little  vesicles 
{Hamburger,^  v.  Harlingen,^  Bruno,^  Ehrmann,^  Brodier  and 
Fouquet,*^  Rawnitzky'').  It  may  happen  that  the  parasite  burrows 
through  a  small  region  of  the  skin  with  many  close  curves  for  some 
time;  on  the  other  hand,  observations  exist  where  it  covered  large 
tracts  in  a  short  time.  The  itching  and  smarting  cease  in  the  place 
left  by  the  larva,  so  that  the  patients  even  in  the  shortest  tract  can 
point  out  at  which  end  the  larva  is,  even  if  they  have  not  watched 
the  lengthening  of  the  tract.  Very  rarely  the  larva  invades  the 
mucous  membrane  of  the  mouth,  the  nose,  and  the  conjunctiva, 
proceeding  from  thence  to  the  external  cutaneous  area. 

The  localization  of  the  affection  is  very  varied  ;  the  primary  seat 
has  been  observed  on  the  glutei  muscles  (Lee,  Kengsep,  Morris,^ 
Rille,  Seifert)  and  their  surroundings  (Stelwagon,  Hamburger, 
Bruno),  on  the  lower  extremities  (Stelwagon,  Lenglet  and  Delaunay, 
Hutchins,    Moorhead,  Lee,  Crocker,  Schmid,^  v.  Harlingen),  on  the 

'  Kengsep,  Derm.  Centralbl.y  1906,  vii. 

2  Hamburger,  y<7«/-«.  of  Cut.  Dis.^  1904. 

'^  V.  Harlingen,  loc.  cit. 

^  Bruno,  v.  Rille  and  Riecke,  "  Handb.  d.  Hautkrankh.  v.  Mracek." 

^  Ehrmann,  loc.  cit. 

^  Brodier  and  Fouquet,  Bull,  de  la  Soc.frauf.  d.  Derm.,  1904. 

'  Rawnitzky,  loc.  cit. 

^  Morris,  Brit.  /onrn.  Derm.,  1896. 

"  Schmid,   Verein  der  Aerzte  in  Steiermark,  February  12,   1900. 


SUPPLEMENT 


731 


upper  extremities  (Samson,  Meade  and  Freeman,  Hutchins,  Sokoloff, 
V.  Harlingen,  Brodier  and  Fouquet,  Shelmire,^  Stelwagon),  on  the 
face  (Sokoloff,  Moorhead,  Kumberg,^  Rawnitzky,  Crocker,  Boas^), 
on  the  neck  (Sokoloff),  and  on  the  body  (Ehrmann,  Brodier  and 
Fouquet,  Kaposi,^  Topsent*''). 

The  duration  of  the  affection  varies  very  much  ;  it  varies  between 
a  few  hours  and  some  years*^ ;  several  times  a  spontaneous  recovery 
has  been  reported. 

The  diagnosis  of  the  disease  is  not  at  all  difficult  owing  to  its 
peculiar  appearance. 

The  treatment  can  only  consist  in  the  removal  or  killing  of  the 
larvae,  since  one  cannot  rely  on  spontaneous  recovery,  even  if  it  has 
occurred  in  some  cases.  If  one  should  succeed  in  locating  the  larva 
as  a  black  spot  at  the  end  of  the  tract,  its  removal  by  means 
of  a  needle  is  the  simplest  method  (Quortrup  and  Boas^).  In 
some  instances  a  cure  has  been  successfully  accomplished  by  excision 
of  the  active  end  of  the  tract  (v.  Neumann  and  Rille,  Schmid).  In 
opposition  to  this  method,  which  not  all  patients  will  allow,  the 
method  practised  by  Arab  women  (Rille  and  Riecke^)  of  killing  the 
worm  with  red  hot  needles  is  quite  rational.  Shelmire^  used  the 
electrolytic  needle  for  the  destruction  of  the  maggots,  Stelwagon^^ 
made  use  of  cataphoresis,  by  means  of  which  he  applied  a  sub- 
limate solution,  afterwards  cauterizing  with  a  drop  of  nitric  acid, 
as  excision  was  refused.  Crocker"  and  v.  Harlingen^^  injected 
small  quantities  of  carbolic  acid;  Moorhead^^  by  a  single  freezing 
of  the  skin  with  ethyl  chloride,  attained  a  definite  cessation  of  the 
attack  at  the  active  end.  Hutchins^^  in  one  case  made  use  of  hypo- 
dermic injection  of  a  few  drops  of  solution  of  cocaine  and  afterwards 
of  I  to  2  drops  of  chloroform  ;  in  a  second  case  of  repeated  applica- 
tions    of    tincture     of    iodide,     as     Lenglet     and     Delaunay^^    did. 


'  Shelm'ne,  /ourn.  Cut.  Dis.,  1905. 

2  Kumberg,  St.  Peter sb.  med.  Wochenschr.,  1898. 

'  Boas,  Monatsh.f.  prakt.  Derm.,  1907,  xliv. 

^  Kaposi,  Wien.  klin.  Wochenschr.^  1898. 

^  Topsent,  Arch,  de  Par.,  1901. 

6   [This  is  extremely  unlikely,  as  the  bots  of  Hypoderma  only  live  for  nine  or  ten  months 
at  the  most !— F.  V.  T.] 

'^  Quortrup  and  Boas,  Hospitalstid.,  1907. 

**  Rille  and  Riecke,  "  Handb.  d.  Hautkrankh.,"  v.  Mracek,  1907,  iv. 

^  Shelmire,  loc.  cit. 
"  Stelwagon,  loc.  cit. 
1^  Crocker,  loc.  cit. 
"  V.  Harlingen,  loc.  cit. 
'^  Moorhead,  Texas  Med.  News,  1906. 
"  Hutchins,  y<7«rw.  Cut.  Dis.,  1906. 
^•^  Lenglet  and  Delaunay,  loc.  cit. 

46 


732  THE    ANIMAL    PARASITES    OP^    MAN 

V.  Harlingen'  allayed  the  affection  in  his  first  case  by  rubbing  in 
sapo  viridis  and  tar,  in  Kensep's^  case  the  cure  seems  to  have  been 
accomplished  by  an  ointment  containing  resorcin,  in  Meade  and 
Freeman's'^  case  by  a  20  per  cent,  ichthyol  paste.  In  our  case  we 
made  exclusive  use  of  Lassar's  paste ;  within  four  weeks  a  cure 
resulted,  probably  spontaneously,  since  one  cannot  ascribe  any 
essential  effect  to  this  paste. 


*  V.  Harlingen,  loc.  cit.  2  Kensep,  loc.  cit. 

^  Meade  and  Freeman,  Brit.  Jourjt.  Deriu.y  October,  1906. 


APPENDIX   ON    PROTOZOOLOGY  733 


APPENDIX  ON  PROTOZOOLOGY, 

Comprising  Notes  on  Recent  Researches,  Formulae  of  some 
Culture  Media,  and  Brief  Notes  on  General  Protozoological 
Technique. 

BY 

H.  B.  FANTHAM,  M.A.,  D.Sc. 


I.— NOTES  ON  RECENT  RESEARCHES. 

Since  the  foregoing  section  on  Protozoology  was  sent  to  press, 
certain  interesting  observations  and  results  have  been  published. 
Brief    notes  on  such,    and  some   references  thereto,  are  now  added. 

It  is  necessary,  however,  to  remark  that  sometimes  it  is  impossible 
to  give  a  precise  or  rigid  definition  to  a  genus  of  Protozoa,  owing  to 
differences  of  opinion,  to  differences  regarding  nomenclature  or  to 
incompleteness  of  knowledge.  Such  a  lack  of  definition,  while  incon- 
venient for  the  time  being,  is  not  unhopeful,  as  it  directs  attention  to 
the  necessity  for  further  work,  which  is  inevitable  in  such  a  relatively 
new  and  wide  subject  as  protozoology.  Thus,  it  may  be  noted  in 
illustration  that  Minchin,  in  1912,  in  his  text-book  regarding  the 
genus  Efitaniceha  writes  :  '^  The  entozoic  amoebae  are  commonly 
placed  in  a  distinct  genus,  Entamoeha,  distinguished  from  the  free- 
living  forms  by  little,  however,  except  their  habitat  and  the  general 
(but  not  invariable)  absence  of  a  contractile  vacuole." 

DiflPerences  between  Entamoeba  histolytica  and  E.  coli. — In 
continuation  of  the  remarks  on  pp.  34  and  40,  it  may  be  added  that 
Lugol's  solution  (iodine  in  aqueous  potassium  iodide  solution)  in 
fresh  specimens  shows  by  brownish  staining  the  presence  of  glycogen 
in  the  vacuoles  of  Entaniceba  coli.  Such  a  reaction  is  rarely  or  never 
given  by  E.  histolytica. 

Phagedaenic  Amoebae. — Carini  and  others  record  cases  in  which 
the  skin  around  an  operation  wound  in  connection  with  liver  abscess 
became  gangrenous.  Amoebae,  possibly  Entamoeba  histolytica,  were 
found  therein  and  may  have  been  responsible  for  the  gangreno- 
phagedaenic  action. 

Endamoeba  ginglvalis  (see  pp.  43,  44). — Smith  and  Barrett,^  after 

*  Joui'TU  of  Parasitol.,  i,  p.  159. 


734  I^HE   ANIMAL   PARASITES   OF   MAN 

analysing  the  early  literature,  state  (June,  191 5)  that  Endamceba 
gingivalis,  Gros,  1849,  is  the  correct  name  for  the  following  organisms  : 
E.  buccalisy  Prowazek,  1904  (see  p.  43)  ;  Amoeba  gingivalis,  Gros,  1849; 
Aniceba  buccalis,  Steinberg,  1862,  and  Amoeba  dentalis,  Grassi,  1879. 
They  conclude  that  E.  gingivalis  is  the  causal  agent  of  pyorrhoea 
alveolaris,  and  that  this  disease  responds  to  treatment  with  emetine. 

Entamoeba  kartulisi  (see  p.  44),  synonym  E.  maxillaris,  Kartulis, 
is  considered  to  be  E.  gingivalis. 

Smith  and  Barrett  adopt  the  generic  name  Endamoeba,  Leidy, 
1879  {see  footnote  on  p.  31,  also  p.  34).  Leidy  worked  on  Endamoeba 
blatta^. 

Cralgia  and  Cralgiasis  {see  p.  45). — Barlow^  (May,  191 5)  found 
Craigia  {Paramoeba)  hominis  in  cases  of  chronic  diarrhoea  and  mild 
dysentery  in  Honduras.  He  also  described  a  new  species  of  Craigia 
under  the  name  of  C.  migrans.  Fifty-six  cases  were  studied,  five  of 
which  were  due  to  Craigia  hominis,  the  remainder  to  C.  migrans. 
In  C.  migrans,  each  flagellate,  on  attaining  full  development,  becomes 
an  amoeba  without  dividing.  Each  amoeba  encysts  and  produces 
a  number  of  flagellates  which  are  somewhat  like  cercomonads.  On 
the  other  hand,  in  C.  hominis  the  flagellate  form  produces,  by  longi- 
tudinal fission,  several  generations  of  flagellates  before  entering  upon 
the  amoebic  stage.  The  cysts  of  C.  migrans  contain  fewer  ^'  swarmers  " 
(flagellul^)  than  those  of  C.  hominis,  but  the  ''swarmers"  are  some- 
what larger,  namely,  5  jjl  instead  of  3  /^  in  diameter.  Further,  there  is 
no  accessory  nuclear  body  in  C.  migrans,  but  its  flagellum  stains 
more  deeply  than  that  of  C.  hominis  and  has  a  peculiar  banded 
iippearance. 

Human  Trichomoniasis  {see  pp.  52-56).— Lynch^  (April  and 
May,  1915),  working  in  Charleston,  seems  to  favour  the  view  that 
the  trichomonads  found  in  the  vagina,  urethra,  mouth,  lungs  and 
alimentary  tract  are  one  and  the  same  organism,  and  that  these 
flagellates  may  further  excite  already  existing  inflammatory  condi- 
tions. He  gives  detailed  histories  of  cases  of  {a)  infection  of  the 
vagina  and  gums,  and  (6)  intestinal  infection  manifested  as  intermit- 
tent attacks  of  diarrhoea.  The  flagellates  were  found  in  catarrhal 
vaginal  discharge,  in  blood-stained  scrapings  from  the  gums  (together 
with  Endamceba  buccalis),  and  in  stools  after  a  purge  of  magnesium 
sulphate.  The  parasites  were  tetratrichomonads  {see  footnote,  p.  53), 
that  is,  each  possessed  four  flagella  anteriorly  as  well  as  an  undulating 
membrane.  Lynch  successfully  infected  rabbits  from  the  cases  and 
from  cultures  of  the  parasite.     Encysted  trichomonads  were  seen  in  a 


Amer.  Journ.  Trop.  Dis.  and  Prevent.  Aled.^  ii,  p.  68o. 
/hid.,  p.  627  ;  New    York  Med.  y^^^rw..  May  I,  1915,  ci,  p. 


APPENDIX   ON   PROTOZOOLOGY 


735 


patient's  stools,  in  rabbits  infected  therefrom  and  in  cultures.  The 
culture  medium  used  was  bouillon  acidified  with  about  0*05  per  cent, 
acetic  acid  and  the  cultures  were  maintained  at  30°  C. 

Trichomonads  occur  in  the  digestive  tracts,  for  example,  the  caeca 
of  rats  and  mice  (fig.  422).  In  man  allied  flagellates  can  occur  in 
similar  situations,  as  well  as  in  other  parts  of  the  intestine. 


Fig.  422. —  Trichomonas  from  csecum  and  gut  of  rat:  «,  nucleus; 
bl,  blepharoplast  ;  y?,  flagella  ;  ax^  axostyle  ;  ///,  undulating  membrane  ; 
b,  line  of  attachment  of  undulating  membrane  to  the  body.  X  2,000 
approx.     (Original.) 

Other  trichomonad-like  organisms  have  been  recently  described 
from  the  faeces  of  man,  more  particularly  from  cases  of  chronic 
dysentery  in  the  tropics.  Derrieu  and  Raynaud^  (July»  1914)^  working 
in  Algeria,  found  a  flagellate  possessing  five  free  flagella  anteriorly 
and  an  undulating  membrane  apparently  lateral.  They  named  the 
parasite  Hexamastix  ardin-delteiliy  but  the  generic  name  Hexaniastix 
is  pre-occupied.  Chatterjee^  (January,  1915),  working  in  India,  found 
probably  the  same  flagellate  and  called  it  Pentatrichomonas  bengalensis. 

Chllomastix  (Tetramitus)  mesnili  (see  p.  57). — Alexeieff^  (1914) 
now  places  the  parasite  originally  called  Macrostoma  mesnili,  by 
Wenyon  (1910),  in  the  genus  Chilomastix,  Alexeieff.  The  differential 
characters  of  the  genera  Tetramitus  and  Chilomastix  are  not  especially 
well  marked.  According  to  Alexeieff,  Tetramitus  is  characterized  by 
four  unequal  flagella  (which  he  figures  anteriorly),  a  ventral  cytostome 
in  the  form  of  a  linear  cleft  and  a  pulsatile  vacuole  in  front  of  the 
anterior  nucleus.      Chilomastix,   according   to   the  same  author,   has 


Bull.  Soc.  Path.  Exot.,  vii,  p.  571. 

Ind.  Med.  Gaz.,  1,  p.  5. 

Zool.  Anzeiger,  xliv,  pp.  203,  206  ;  and  ibid.^  xxxix,  p.  678. 


736 


THE   ANIMAL   PARASITES   OF   MAN 


three  forwardly  directed  flagella  and  a  fourth  backwardly  directed  one 
in  the  cytostome,  which  is  well  developed  (fig.  423).  Some  authors 
consider  that  the  fourth  flagellum  forms  the  edge  of  an  undulating 
membrane  in  the  cytostome. 

Diagrams  of  Chilomastix  niesnili  are  given  in  fig.  423. 


Fig.  423. —  Chilomastix  ( 1  etramitus)  mesnili.     a,  b,  c,  flagellate  forms ; 
dy  rounded  or  encysted  form.      X  2,500.     (Original.) 


Giardla  (Lamblia)  intestinalis  {see  p.  57). — Alexeiefif^  (1914)  con- 
siders that  Lamblia  intestinalis,  h^Lmbl,  should  be  placed  in  the  genus 
Giardla,  Kunstler,  1882.  Bipartition  occurs  in  the  encysted  state. 
The  axostyles  persist  in  the  quadrinucleate  cyst. 

Cercomonas  hominis  {see  p.  61). — This  parasite  is  considered  by 
some  authors  to  be  of  a  doubtful  nature,  as  it  is  thought  to  have  been 
mistaken  for  deformed  or  incompletely  observed  Trichomonas  or  Chilo- 
mastix or  even  Lamblia. 

Wenyon^  (1910)  described  Cercomonas  longicauda  from  cultures 
of  human  faeces.  It  is  considered  that  the  genus  is  very  confused, 
and  the  author  points  out  that  the  tail  flagellum  has  been  overlooked. 
He  considers  that  the  genus  Cercomonas  should  include  flagellates 
with  an  anterior  blunt  end  from  which  arises  a  single  long  flagellum, 
and  a  posterior  tapering  end  also  with  a  flagellum,  which  can  be  traced 


'  ZooL  Anzeiger,  xliv,  p.  210. 

-  Quart.  Journ.  Micros.  Sci.,  Iv,  p.  241. 


APPENDIX   ON   PROTOZOOLOGY  737 

over  the  surface  of  the  body  towards  the  insertion  of  the  anterior 
flagellum. 

Another  species,  Ccrcojiionas  parva,  has  been  found  in  cultures  of 
human  faeces  by  Hartmann  and  Chagas^  (1910).  It  has  a  somewhat 
different  structure. 

Furthur  researches  are  necessary  on  the  organisms  variously 
referred  to  the  genus  Cerconwnas. 

Transmlssive  Phase  of  Trypanosomes  in  Vertebrates. — In 
addition  to  the  general  remarks  on  the  morphology  of  trypanosomes 
set  forth  on  pp.  70  to  72,  it  may  be  noted  that  Woodcock^  (November, 

1 9 14)  states  that,  in  certain  cases,  there  is  a  definite  transmissive  phase  of 
a  trypanosome  in  its  vertebrate  host.  He  quotes  the  work  of  Minchin 
and  himself  on  T.  noctiice  of  the  little  owl,  in  which  the  transmissive 
form  is  spindle-shaped  and  occurs  in  the  bird's  peripheral  blood 
during  the  early  summer  months  {see  p.  69).  A  similar  phase  occurs 
in  T.  Jringlllarum,  and  Robertson^  has  found  that  the  short,  stumpy 
form  of  T.  gauibiense  is  its  transmissive  phase  in  vertebrates. 

Trypanosoma  lewisi  {see  p.  88). — Brown  (1914-15)  has  published 
some  interesting  results  on  the  potential  pathogenicity  of  T.  lewisi, 
Blepharoplastless  Trypanosomes  {see  p.  loi). — Laveran^  (April, 

19 1 5)  suggested  a  practical  use  of  strains  of  blepharoplastless  trypano- 
somes produced  by  the  action  of  drugs.  He  finds  that  tryposafrol 
will  also  produce  such  strains,  and  remarks  on  blepharoplastless 
strains  of  T.  evansi  and  T.  brucei,  which  in  the  former  case  can 
undergo  450  passages  without  reversion,  and  in  the  latter  273 
passages.  He  states  that  if  it  is  desired  to  inoculate  surra  or  nagana 
to  Capridae  or  Bovidae  in  order  to  produce  immunity,  use  should  be 
made  of  the  blepharoplastless  races  of  the  respective  trypanosomes, 
which  races  are  a  little  less  virulent  than  the  corresponding  normal 
ones.  Also,  the  immunity  which  follows  from  an  infection  due  to 
blepharoplastless  T.  evansi  or  T.  brucei  is  only  a  little  less  coifiplete 
than  that  following  infections  from  either  of  the  respective  normal 
strains. 

The  Experimental  Introduction  of  certain  Insect  Flagellates 
into  various  Vertebrates,  and  Its  bearing  on  the  Evolution  of 
Leishmaniasis. — In  continuation  of  the  remarks  on  pp.  103,  104, 
and  112,  further  researches  have  been  conducted  on  the  introduction 
into  vertebrates  of  flagellates  normally  parasitic  in  insects.  The 
vertebrates  became  infected  by  inoculation  with  the  flagellates  or  by 


'  A/em.  Inst.  Oswalao  Cruz,  ii,  p.  67. 
^Arch.f.  Protistenk.^  xxxv,  p.  197. 
^  Proc.  Roy.  Soc,  B,  Ixxxv,  p.  527. 
4  C.  R.  Acad.  Set.,  clx,  p.  543. 


738  THE   ANIMAL   PARASITES   OF   MAN 

being  fed  on  insects  containing  the  protozoa.  Fantham  and  Porter^ 
(June,  1915)  published  the  following  results.  Flagellates  from 
sanguivorous  and  non-sanguivorous  insects  were  used,  and  cold- 
blooded as  well  as  w-arm-blooded  vertebrates  as  hosts.  The  intro- 
duced protozoa  were  pathogenic  to  the  mammals,  but  not  markedly 
so  to  the  cold-blooded  vertebrates.  Herpetomonas  jaaihiiu,  H.  stratio- 
myice^  H.  pediaili,  and  Crithidia  gerridis  (parasitic  in  certain  water- 
bugs)  proved  pathogenic  to  mice.  A  puppy  was  infected  by  way 
of  the  digestive  tract  with  H.  ctenocephali.  Frogs  became  infected 
with  H.  jaciilum  and  with  C.  gerridis^  toads  and  grass  snakes  w'lih. 
H.  jacnlum,  lizards  with  C.  gerridis,  and  sticklebacks  with  //.  jaadiim. 
Second  and  third  passages  of  some  of  the  parasites  were  obtained. 
The  protozoa,  whether  Herpetofuonas  or  Crithidia,  were  present  in 
the  vertebrate  hosts  in  either  the  non-flagellate  or  the  flagellate  form, 
or  usually  both.  They  were  more  abundant  in  the  internal  organs 
of  the  hosts,  more  particularly  in  the  liver,  spleen  and  bone-marrow'. 
In  all  experiments  in  which  C.  gerridis  was  used  the  parasite 
invariably  retained  the  crithidial  facies  in  the  vertebrate  host.  No 
transition  to  a  trypanosome  was  ever  seen.  Infections  in  adult 
animals  were  not  so  heavy  as  in  the  young  ones,  and  the  parasites 
were  more  virulent  in  young  hosts,  as  is  the  case  with  Mediterranean 
kala-azar  in  children. 

The  mode  of  infection  of  the  vertebrate  in  Nature  seems  to  be 
contaminative,  either  by  its  food  or  through  an  already  existing 
abrasion  or  puncture  on  the  surface  of  its  body.  Cases  in  which  the 
flagellate-infected  insects  have  been  allowed  to  suck  the  blood  of 
vertebrates  have  proved  negative  up  to  the  present.  In  areas  where 
leishmaniases  are  endemic,  an  examination  should  be  made  of  all 
insects  and  other  invertebrates  likely  to  come  into  contact  with  men 
or  dogs,  or  rats  and  mice  (see  below),  in  order  to  ascertain  if  these 
invertebrates  harbour  herpetomonads.  Preventive  measures  should 
be  directed  against  such  invertebrates,  especially  arthropods.  Further, 
it  is  likely  that  certain  vertebrates,  such  as  reptiles  and  amphibia 
(especially  those  that  are  insectivorous),  may  serve  as  reservoirs 
of  leishmaniases,  or,  as  they  should  preferably  be  termed,  herpeto- 
moniases.  From  such  reservoirs  the  herpetomonads  may  reach 
man  by  the  agency  of  ectoparasites  or  flies,  especially  such  as  are 
sanguivorous. 

That  vertebrates  in  Nature  can  harbour  herpetomonads  in  their 
blood  has  been  shown  by  the  work  of  Button  and  Todd  (1903)  on 
the  herpetomonads  of  Gambian  mice,  while  the  recently  published 


'  Proc.   Camb,  Philosoph.  Soc,  xviii,  p.  137  ;  and  ^nna/s  Trop.  Med.  and  ParasitoL,  ix, 
P-  335- 


APPENDIX   ON   PROTOZOOLOGY  739 

investigations  of  Fantham  and  Porter^  (June,  1915)  on  natural  herpe- 
tomonads  in  the  blood  of  mice  in  England  have  shown  that  these 
rodents  may  be  a  natural  reservoir  of  herpetomoniasis.  The  origin 
of  the  infection  of  mice  is  to  be  sought  in  a  flagellate  of  an  ecto- 
parasite of  the  mouse,  very  probably  Herpetomonas  pattoni  parasitic  in 
various  fleas,  which  protozoon  can  adapt  itself  to  life  in  the  blood 
of  mice.  Herpetomonads  were  also  found  naturally  in  the  blood 
of  birds  by  Sergent  (1907).  Recently,  Fantham  and  Porter  have 
successfully  infected  birds  with  herpetomonads  experimentally. 

The  significance  of  insect  flagellates  in  relation  to  the  evolution  of 
disease  has  recently  been  set  forth  by  Fantham^  (June,  1915).  The 
deductions  to  be  made  from  the  occurrence  of  a  herpetomonad  stage 
in  Leishmania,  especially  in  L.  tropica,  in  man  himself,  and  of  flagel- 
late stages  of  the  so-called  Histoplasma  capsiilainm  in  man  are  fully 
discussed  and  correlated.  It  is  pointed  out  that  flagellosis  of  plants 
(see  p.  104)  may  possibly  be  connected  with  leishmaniasis.  The 
evolution  of  Lcishmanla  from  flagellates  of  invertebrates  is  thus 
traced  and  the  way  again  indicated  for  preventive  measures  against 
leishmaniasis,  as  first  set  forth  by  Dodds  Price  and  Rogers. 

Franchini  and  Mantovani  (March,  191 5)  have  successfully 
infected  rats  and  mice  by  inoculation  or  by  feeding  with  Herpeto- 
monas muscat  domestical  obtained  from  flies  and  from  cultures. 

It  is  of  great  interest  to  note  that  the  recent  observations  of 
Ed.  and  Et.  Sergent,  Lemaire  and  Senevet^  (19 14)  have  demonstrated 
the  presence  of  a  herpetomonad  flagellate  in  cultures  of  the  blood 
and  organs  of  geckos  obtained  from  areas  in  Algeria  in  which  Oriental 
sore,  due  to  L.  tropica,  is  prevalent.  Phlebotomiis  flies,  which  may 
harbour  a  natural  herpetomonad,  feed  on  the  geckos  and  on  men. 
Hence  animals  like  geckos  may  possibly  act  as  reservoirs  of 
leishmaniasis.  Lindsay'^  (1914)  writes  that  the  parasite  of  dermo- 
mucosal  leishmaniasis  in  Paraguay  is  believed  by  native  sufferers  to 
be  conserved  in  rattlesnakes,  and  spread  by  ticks  or  flies  {Simutium) 
feeding  on  the  reptiles  and  transferring  the  parasite  to  man. 

The  Transmission  of  Spirochaeta  duttoni  (see  p.  116). — It  is 
probable  that  Ornitlwdorus  savignyi  acts  as  the  transmitting  agent  of 
S.  c/////o;//  in  places  like  Somaliland  (Drake-Brockman,  1915).'' 

Spirochaeta  bronchialis  {see  p.  122). — The  morphology  and  life- 
history  of  S.  broncliialis  have  been  investigated  by  Fantham^  (July, 
1915).     From  researches  conducted  in  the  Anglo-Egyptian  Sudan,  he 

'  Parasitology,  viii,  p,  128. 

-  Annals  Trop.  Med.  and  FarasiloL,  ix,  p.  335. 

=*  Bull.  Soc.  Path.  Exot.,  vii,  p.  577. 

4  Trans.  Soc.  Trop.  Med.  and  Hyg.,  vii,  p.  259. 

^  Ibid.,  viii,  p.  201. 

^  Annals  Trop.  Med.  and  Parasitol.,  ix,  p.  391. 


740  THE    ANIMAL    PARASITES    OF   MAN 

found  that  S.  bronchialis  is  an  organism  presenting  marked  poly- 
morphism, a  feature  that  has  only  been  determined  by  the  examination 
of  numerous  preparations  from  the  deeper  bronchial  regions  of  various 
patients. 

S.  bronchialis  varies  in  length  from  5  ft  to  27  fi,  and  its  breadth  is 
about  0-2/A  to  o-6yLt.  These  variations  are  due  to  the  processes  of 
growth  and  division.  Many  of  the  parasites  measure  either  14  yu-  to 
16 />t  long,  or  7 /A  to  9yLt,  the  latter  resulting  from  transverse  division 
of  the  former.  The  ends  show  much  variation  in  form,  but  approach 
the  acuminate  type  on  the  whole.  The  discrepancies  in  dimensions 
given  by  the  very  few  previous  workers  on  the  subject  are  probably 
the  result  of  the  measurement  of  a  limited  number  of  parasites.  All 
such  sizes  can  be  found  on  some  occasion  during  the  progress  of  the 
disease,  when  a  larger  number  of  spirochsetes  is  examined. 

The  movements  of  S.  bronchialis  are  active,  but  of  relatively  short 
duration,  when  it  is  removed  from  the  body.  The  number  of  coils 
of  the  spirochaete  is  rather  an  index  of  its  rapidity  of  motion  than  a 
fixed  characteristic  of  the  species. 

The  motile  phase  of  S.  bronchialis  is  succeeded  by  one  of  granule 
formation,  the  granules  or  coccoid  bodies  serving  as  a  resting  stage 
from  which  new  spirochaetes  are  produced.  The  formation  of 
coccoid  bodies  and  reproduction  of  spirochaetes  from  them  can  be 
observed  in  life. 

S.  bronchialis  is  a  species  distinct  from  the  spirochaetes  occurring 
in  the  mouth.  It  differs  from  them  in  morphology,  pathogenicity 
and  in  staining  reactions.  It  is  not  a  developmental  form  of  any 
bacterium,  and  is  an  entity  in  itself. 

The  passage  from  man  to  man  is  effected  most  probably  by  means 
of  spirochaetes,  and  especially  coccoid  bodies,  that  leave  the  body  in 
the  spray  with  expired  air  and  byway  of  the  nasal  secretions.  Owing 
to  the  fragility  and  short  life  of  S.  bronchialis  extracorporeally,  the 
resistant  coccoid  bodies  in  air,  in  dried  sputum  and  dust,  and  possibly 
also  on  the  bodies  of  flies  and  other  insects,  are  probably  instrumental 
in  inducing  attacks  of  bronchial  spirochaetosis  in  human  beings,  espe- 
cially those  having  a  lowered  bodily  resistance,  such  as  occurs  after 
a  chill.     Lurie  (December,  1915),  has  described  a  case  from  Serbia. 

The  Spirochaetes  of  the  Human  Mouth  {see  p.  122). — Two 
species  of  spirochaetes  were  recorded  as  occurring  in  the  human 
mouth  about  forty  or  fifty  years  ago.  These  are  Spirochccta  buccalis, 
Steinberg  (often  ascribed  to  Cohn,  1875),  and  S.  dentinni,  Miller 
(often  attributed  to  Koch,  1877). 

The  most  recent  work  on  S.  dentinm  and  S.  bnccalis  is  that  of 
Fantham^  (July,  1915),  who  observed  the  parasites  ascribed  to  Cohn 

*  Annals  'Irop.  Med.  and  ParasitoL,  ix,  p.  402. 


APPENDIX  OX  PROTOZOOLOGY  74I 

and  to  Koch,  these  being  the  two  common  spirochcetes  seen  in  the 
mouths  of  natives  of  the  Sudan  and  of  Europeans  in  England,  as 
well  as  the  forms  described  and  cultivated  by  recent  investigators. 
Some  of  the  mouth  spirochaetes  are  not  very  active,  but  there  is 
marked  corkscrew  and  boring  movement,  and  they  are  flexible. 
Tangles  or  tomenta  of  these  mouth  spirochetes  are  common. 
Internal  structure  is  seen  with  some  difficulty,  but  in  some  specimens 
it  can  be  determined,  and  chromatin  granules  are  then  seen.  Muhlens 
(1907)  figured  stained  specimens  of  S.  hiiccalis  and  S.  dentiiim,  in 
which  chromatin-coloured  granules  were  distributed  along  the  bodies 
of  the  organisms. 

S.  deiitiiim  has  tapering  ends,  and  varies  in  length  from  4  //.  to  lOyLt. 
S.  dentitun  is  rather  like  Treponema  pallidum,  and  has  been  placed  by 
some  workers— for  example,  Dobell— in  the  genus  Treponema.  It  has 
already  been  mentioned,  on  p.  128,  that  Noguchi  cultivated  three 
species  of  Treponema  from  the  human  mouth — namely,  T.  macro- 
denthun,  T.  microdentium,  and  T.  mucosum,  but  they  cannot  be  easily 
distinguished  morphologically,  and  so  may  appear  to  be  biological 
varieties  of  S.  dentium. 

S.  huccalls  has  somewhat  rounded  or  bluntly  acuminate  ends 
and  varies  in  length  from  g  fi  to  22 /jl.  A  slight  membrane  or  crest 
may  sometimes  be  observed.  S.  huccalls  was  found  to  be  the  pre- 
dominant spirochaste  in  the  mouths  of  eight  natives  examined  by 
Fantham  in  the  Anglo-Egyptian  Sudan. 

S.  huccalls  and  S.  dentium  take  up  stains  well  and  with  relative 
ease.  Intracellular  stages  of  the  parasites  are  uncommon.  Multiplica- 
tion by  binary  fission  has  also  been  observed.  Coccoid  bodies  or 
granule  stages  of  the  mouth  spirochaetes  are  formed,  but  appear  to  be 
relatively  few  in  number. 

J.  G.  and  D.  Thomson^  {1914)  have  written  an  interesting  paper 
on  various  spirochaetes  occurring  in  the  alimentary  tract  of  man  and 
of  some  of  the  lower  animals.  They  have  also  given  a  useful  list  of 
references,  and  the  work  of  some  of  the  earlier  authors  is  discussed  in 
the  paper. 

With  regard  to  the  general  morphology  of  spirochaetes,  it  may  be 
noted  that  the  so-called  axial  fibre  of  Zuelzer  is  acknowledged  to  be 
homologous  with  the  membrane  or  crista  of  molluscan  spirochaetes. 

Coccidia  in  Cattle. — Regarding  the  remarks  on  coccidiosis  or 
**  red  dysentery  "  in  cattle  on  p.  147,  it  may  be  added  that  Schultz^ 
(July,  1915)  has  found  the  malady  among  cattle  in  the  Philippine 
Islands.  He  states  that  some  irregular  or  atypical  cases  of  apparent 
rinderpest  are  really  due  to  coccidia.     As  has  been  pointed  out  by 


Proc.  Roy.  Soc.  Med.^  vii,  pt.  i,  p.  47.  "  Jotirn.  Infect.  Dis.,  xvii,  p.  95. 


742  THE   ANIMAL   PARASITES   OF   MAN 

Montgomery,  rinderpest  can  be  transmitted  by  blood  inoculation, 
while  coccidiosis  cannot  be  so  transmitted,  but  may  be  diagnosed  by 
the  microscope.  These  differences  should  be  remembered  as  the  two 
diseases  are  often  found  to  be  associated  and  are  difHcult  to  separate 
clinically.     Coccidia  have  also  been  found  in  Australian  cattle. 

The  Haemosporidia. — It  is  likely  that  this  order  {see  p.  151)  may 
be  soon  abolished.  MesniP  (April,  19 15)  considers  that  the  grouping 
of  the  three  families,  Plasmodiidae  (or  Haemamcebidae),  Haemogreg- 
arinidae  and  Piroplasmidae  in  the  order  Haemosporidia  is  no  longer 
possible,  because  of  the  coccidian  nature  of  the  Haemogregarines 
{see  p.  154).  The  Coccidia  are  divisible  into  the  Adeleidea  and  the 
Eimeridea  (see  p.  141).  The  Haemogregarinidae  are  allied  to  the 
former,  and  the  Plasmodiidae  to  the  latter.  The  Piroplasmidae,  until 
more  is  known  of  their  life-cycle  in  the  invertebrate  host,  cannot  be 
more  definitely  placed. 

The  Leucocytozoa  of  Birds. — Regarding  the  statement,  on  p.  153, 
that  Laveran  and  FYanca  consider  that  avian  leucocytozoa  may  inhabit 
red  blood  cells,  it  may  be  added  that  Franga''^  (April,  1915)  remarks 
that  the  action  of  the  parasites  on  the  red  cells  is  very  rapid  and 
very  intense.  The  host  cells  become  so  altered  that  it  is  difficult 
to  -recognize  their  true  nature.  He  used  very  young  birds  in  his 
researches.  Two  shapes  of  host  cell  are  considered,  namely,  those 
with  fusiform  prolongations,  and  those  which  are  rounded  and  with- 
out such  prolongations  (see  p.  153).  The  movements  and  form  of 
the  Leucocytozoa  determine  the  shape  of  the  host  cell,  as  was  pointed 
out  by  Fantham^  in    1910. 

Schizogony  of  these  parasites  has  been  seen  by  Franga  (191 5)  and 
by  Coles  (1914),  in  addition  to  Fantham  (1910),  and  to  Moldovan 
(1913),  mentioned  on  p.  153.  Schizogony  may  also  take  place  in  the 
lungs  of  the  host.  The  genus  Leucocyiozodn,  established  by  Ziemann 
in   1898,  belongs  to  the  family  Haemamoebidae. 


II.— FORMUL/E  OF  SOME  CULTURE  MEDIA. 

(i)  Culture  Media  for  growing  Amoebae.— There  has  been  much 
discussion  as  to  whether  the  true  parasitic  Entamcebcv  or  Endamcehce 
can  be  grown  on  culture  media  [see  p.  42).  Undoubtedly  certain 
free-living  amoebae  can  be  so  grown,  and  it  is  considered  that  some 
of  the  earlier  researches  on  the  so-called  artificial  growth  of  the 
dysenteric  amoebae  were  really  due  to  contaminations  with  free-living 
forms.     The  following  media  are  worthy  of  note  : — 

'  Bid  I.  Soc.  Path.  Exot.,  viii,  p.  241. 

-  Ibid.,  p.  229.  3  Pj.^^^  2:001.  Soc.  Lond.,  1910,  p.  694 


APPENDIX  ON  PROTOZOOLOGY  743 

Musgrave  and  Clegg  in  1904  devised  a  culture  medium  for 
amoebae.  The  organisms  grown  by  them  were  probably  not  dysen- 
teric amoebae,  as  was  thought,  but  free-living  forms.  Phillips^  (1915) 
•gives  a  slightly  modified  formula  of  Musgrave  and  Clegg's  medium, 
thus : — 

Agar-agar                  ...  ...  ...  ...  2*5    grm. 

Sodium  chloride      ...  ...  ...  ...  0*05     ,, 

Liebig's  beef  extract  ...  ...  ...  0*05     ,, 

Normal  sodium  hydroxide  ...  ...  ...  2*0     c.c. 

Distilled  water        ...  ...  ...  ...  100*0       ,, 

Without  clarifying,  sterilize  at  7  kilograms  pressure  per  square  centi- 
metre for  about  three-quarters  of  an  hour.  It  should  be  neutral  to 
phenolphthalein. 

Anna  W.  Williams"  (191 1)  described  a  medium  consisting  of  fresh 
tissue  spread  on  agar  plates  for  the  culture  of  amoebae.  There  are 
three  stages  in  the  procedure  :  (i)  obtaining  living  amoebae  free  from 
other  living  organisms ;  (2)  obtaining  sterile  tissue  ;  and  (3)  making 
successive  transplants  of  amoebae  and  tissue,  and  showing  that  every 
transplant  is  free  from  other  living  organisms.  Each  step  requires 
many  controls.  The  essentials  of  the  method  may  now  be  given. 
Remove  aseptically  and  rapidly  the  tissue  required,  such  as  brain, 
liver,  kidney,  or  spleen,  from  a  freshly  killed  animal  (guinea-pig, 
rabbit,  or  dog).  Put  each  tissue  on  a  separate  agar  plate.  Cut  the 
selected  tissue  into  tiny  pieces,  and  spread  them  over  freshly  made 
agar  plates.  Place  these  plates  in  a  thermostat  at  36°  C.  for  tw^enty- 
four  hours  to  insure  sterility.  Add  the  broken  up  tissue  to  the 
amoebae,  free  from  bacteria,  and  maintain  the  cultures  in  thermostats, 
some  at  36°  C,  and  some  at  20°  C.  to  24°  C.  Emulsions  of  liver  and 
brain  in  sterile  neutral  glycerine  may  also  be  used.  The  freshly 
removed  tissue  serves  as  food  for  the  amoebae. 

The  cultural  amoebae  mentioned  on  p.  42  were  grown  on  such 
media  or  modifications  thereof.  One  modified  medium  actually  used 
was  brain  tissue,  to  which  blood  was  added  from  day  to  day,  and  an 
easily  assimilable  bacterium  (one  of  the  influenza  group  of  bacilli) 
was  present,  which  did  not  overgrow  the  medium  at  a  temperature  of 
38°  C.  Different  conditions  of  food  and  of  temperature  produced 
morphological  variations  in  the  cultural  amoebae. 

Couret  and  ].  Walker^  (1913)  state  that  they  have  cultivated  five 
varieties  of  intestinal  amoebae,  the  associated  bacteria  having  been 
previously  separated.     They  used   a  medium  consisting  of  agar   to 


'  *'  Amoebiasis  and  the  Dysenteries,"  p.  8. 

-  Journ.  Med.  Research,  xxv,  p.  263  ;  and  Proc.  Soc.  Exper.  Biol,  and  Med.,  viii,  p.  56. 

3  Tourn.  Exper.  Med.,  xviii,  p.  252. 


744  THE   ANIMAL   PARASITES   OF   MAN 

which  sterile  autolysed  tissue  had  been  added.  The  sterile  tissue, 
such  as  brain  or  liver,  was  kept  in  a  sterile  thermostat  at  a  temperature 
of  40°  C.  for  ten  to  twenty  days.  The  surface  of  the  agar  should 
be  broken  up  before  use,  and  the  medium  must  not  be  too  acid  (not 
over  I '5  per  cent.).  They  consider  that  autolysed  tissue  is  necessary 
for  the  growth  of  Entamoebae,  and  that  naturally  associated  bacteria 
aid  growth  by  autolysing  the  tissues. 

(2)  Culture  Media  for  the  growth  of  Protozoa  parasitic  in  the 
Blood. — MacNeal  and  Novy,^  in  1903,  used  a  mixture  of  blood  and 
agar  for  the  cultivation  of  trypanosomes  such  as  T.  lewisi  and  T.  brucei. 
They  employed  varying  proportions  of  the  blood  and  agar,  a  medium 
consisting  of  two  parts  of  deiibrinated  rabbit's  blood  mixed  with  one 
part  of  agar  being  useful.  The  trypanosomes  grew  in  the  water  of 
condensation.  Some  of  the  authors'  earlier  formulae  contained 
different  proportions  of  blood  and  agar  w^ith  a  little  peptone,  while 
one  of  these  media  contained  meat  extract,  agar,  peptone,  salt  and 
sodium  carbonate.  The  temperature,  like  the  proportion  of  blood 
and  agar,  varied  with  the  trypanosome  investigated,  but  the  optimum 
was  25°  C. 

Mathis^  (1906)  somewhat  simplified  the  technique  of  Novy  and 
MacNeal.  He  collected  the  blood  of  a  suitable  animal,  such  as  rabbit, 
cow  or  dog,  strict  asepsis  not  being  essential.  The  blood  was 
defibrinated  in  the  ordinary  way.  One  part  of  blood  was  added 
to  two  parts  of  agar  at  50°  C.  The  mixture  was  sterilized  several 
times  by  heating  to  75°  C.  or  100°  C.  Slopes  were  made  and  the 
water  of  condensation  was  inoculated  with  a  little  blood  containing 
the  trypanosomes.  Blood  may  be  obtained  from  a  superficial  vein 
or  from  the  heart. 

Novy-MacNeal-Nicolle  or  N.  N-.  N.  Medium. — In  1908  C.  Nicolle^ 
brought  forward  a  modification  of  the  Novy-MacNeal  (N.N.) 
medium.     The   formula    is   as   follows  : — 

Agar  ...  ...  ;..  ...  ...  I4grni. 

Sea  salt     ...  ...  ...  ...  ...  6     ,, 

Water        ...  ...  ...  ...  ...       900     ,, 

Apparently  pure  sodium  chloride  can  be  substituted  equally 
well  for  sea  salt.  The  mixture  is  placed  in  tubes  and  sterilized  in 
an  autoclave.  To  each  tube  one-third  of  its  volume  of  rabbit 
blood,  taken  by  aseptic  puncture  of  the  heart,  is  added.  The  salt 
agar  is  kept  liquid  at  45°  C.  to  50°  C.  and  the  blood  is  added  to 
the  mixture.  The  culture  medium  so  prepared  is  maintained  for 
five    days   at  37°  C,  and  then  for  a  few  days  at  room  temperature. 


'  See  Sleeping  Sickness  Bulletin  (1909),  i,  No.  8,  p.  287. 

2  C.  R.  Soc.  Biol,  Ixi,  p.  550.  3  c^  ^,  ^^^^  ^^i^  (,jj1^^  p_  g^^. 


APPENDIX   ON    PROTOZOOLOGY  745 

This  medium  was  devised  for  the  cultivation  of  Leishmania  {see 
p.  106),  but  trypanosomes  may  also  be  grown  thereon.  Subsequently, 
NicoUe  recommended  the  use  of  citrated  rat's  blood  heated  to  45°  C. 
for  half  an  hour,  instead  of  defibrinated  rabbit's  blood.  On  such  a 
medium,  J.  G.  Thomson  and  Sinton^  (19 12)  succeeded  in  growing 
Trypanosoma  ganihiense  and  T.  rhodesieiise  (see  pp.  76,  83). 

Noguchi's  media  for  the  cultivation  of  Spirochaetes  and  Trepo- 
nemata  are  described  on  pp.  123,  125.  Hata's  modification  is 
discussed   on    p.    126. 

Bass's  glucose-blood  medium  for  the  cultivation  of  malarial 
parasites  is  described  on  pp.  170-172.  It  has  also  been  used 
successfully  for  the   cultivation  of  Piroplasnia  or  Babesia  {see  p.  172). 

III.— BRIEF  NOTES  ON  GENERAL  PROTOZOOLOGICAL 

TECHNIQUE. 

The  object  of  this  book  is  to  give  accounts  of  the  structure  and 
life-histories  of  the  numerous  parasitic  organisms  that  affect  man 
more  particularly.  It  is,  therefore,  inappropriate  to  devote  much 
space  to  a  consideration  of  technique,  regarding  which  many  volumes 
have  already  been  written.  Methods  of  procedure  are  largely  matters 
of  opinion,  and  the  technique  that  gives  brilliant  results  when  used 
by  one  investigator  may  be  a  complete  failure  in  the  hands  of 
another.  In  the  present  appendix,  brief  notes  regarding  certain 
relatively  simple  methods  only  can  be  given,  because  the  number  of 
fixatives  in  use  is  very  great ;  there  are  also  large  numbers  of  stains 
as  well  as  many  modifications  of  them,  while  the  methods  of  apply- 
ing both  fixatives  and  stains  are,  perhaps,  still  more  numerous. 
There  are  so  many,  in  fact,  that  confusion  frequently  arises  from 
the  multiplicity  of  choice  presented  to  the  worker.  Those  desiring 
more  information  on  the  subject  of  technique  are  advised  to  consult 
the  treatises  of  Bolles  Lee^  and  of  Langeron.-^ 

Fresh  Material. 

{a)  Simple  Examination. 

Fluid  Substances,  such  as  Blood  and  Sputum. — A  small  quantity  of 
the  substance  to  be  examined  is  taken  on  a  sterile  platinum  loop  and 
transferred  to  a  perfectly  clean  glass  slide.  A  clean  cover-slip  is  gently 
lowered  on  to  the  drop,  air  bubbles  being  avoided.     The  preparation 

^  Annals  Trop.  Med.  and  ParasitoL,  vi,  p.  331. 

*  "  The  Microtomist's  Vade  Mecum  "  (7th  edition,  1913).    London  :  J.  and  A.  Churchill. 

^  '*  Precis  de  Microscopic"  (1913).     Paris  :  Masson  et  Cie. 


746  .  THE   ANIMAL   PARASITES   OF   MAN 

is  luted  with  vaseline  or  paraffin  and  examined  first  with  a  low  power 
and  then  with  a  high  power  objective.  The  light  is  cut  down  by 
partly  closing  the  diaphragm  of  the  substage  of  the  microscope. 

Skin  Ulcers  and  Similar  Sores. — Scrapings  are  made  from  the  edge 
of  the  sore,  mixed  with  sterile  physiological  salt  solution,  and 
prepared  and  examined  as  above. 

Fceces. — A  small  portion  of  faeces,  or  flakes  of  mucus  (which  may 
be  blood-stained)  from  the  same,  is  removed  on  a  sterile  platinum 
loop,  spread  out  thinly  after  dilution,  if  necessary,  with  physiological 
salt  solution  on  a  slide,  covered  and  examined  as  before. 

Alternatively,  hanging  drop  preparations  of  blood,  ulcerative  tissue, 
or  faeces,  appropriately  diluted  if  necessary  with  sodium  citrate  or 
physiological  salt  solution,  may  be  made  on  a  cover-slip,  which  is 
inverted  over  a  slide  with  a  well  in  it.  The  cover-slip  is  then  luted  and 
examined. 

For  the  elucidation  of  the  developmental  processes  of  such 
organisms  as  trypanosomes,  spirochaetes  and  piroplasms,  fresh 
preparations  may  be  often  kept  under  observation  longer  by  the 
use  of  a  thermostat,  maintained  at  or  near  blood  heat,  in  which  the 
microscope  is  inserted. 

(b)  Intra  vitaui  Staining  of  fresh  Preparations. 

intra  vitani  staining  is  of  service  on  some  occasions,  more 
particularly  for  the  study  of  the  nucleus  and  other  chromatoid 
substances  of  the  living  organism.  Two  methods  are  in  common 
use.  In  the  first  case,  the  stain,  employed  usually  in  very  dilute 
solution,  is  mixed  wdth  the  medium  containing  the  organism.  The 
latter  takes  up  some  of  the  stain,  the  amount  of  coloration  depending 
on  the  organism  concerned  and  on  the  stain  employed. 

The  commoner  intra  vitam  stains  are  pure,  medicinal  (zinc-free) 
methylene  blue  and  neutral  red,  used  in  aqueous  solutions.  A  solution 
of  methylene  blue  of  i  per  1,000  of  water  may  be  tried,  while  neutral 
red  in  the  proportion  of  i  per  3,000  parts  of  water  has  proved  of 
service. 

The  second  method  of  vital  colouring  consists  in  placing  a  drop  of 
I  per  cent,  solution  of  methylene  blue  on  a  slide  or  cover-slip,  slightly 
spreading  it,  and  allowing  it  to  dry.  The  living  organism  is  then 
placed  in  a  drop  of  saline  on  the  prepared  slide  or  cover-slip,  which 
is  then  mounted  and  examined  under  the  microscope.  Progressive 
staining  of  the  organism  occurs  and  its  internal  structure  can  be  seen. 
A  similar  procedure  may  be  followed  for  neutral  red.  Intra  vitam 
staining  is  useful  for  relatively  large  and  easily  deformed  protozoa 
such  as  ciliates,  as  well  as  for  amoebae  and  flagellata  of  the  gut. 

When  examining  very  actively  motile  organisms,  it  is  sometimes 


APPENDIX  ON  PROTOZOOLOGY  747 

useful  to  endeavour  to  restrict  their  movements  by  adding  a  little  gum 
or  gelatine  to  the  medium. 

(c)  Examination  by  aid  of  the  Paraboloid  Condenser, 

The  use  of  one  of  the  dark-ground  illuminators  (so-called  ultra- 
microscopes)  is  of  service  for  the  detection  of  minute  living  organisms 
or  of  organisms  present  in  small  numbers  only.  The  forms  of 
paraboloid  condenser  manufactured  by  the  firms  of  Zeiss  and  Leitz 
can  be  recommended.  For  details  of  their  methods  of  employment, 
reference  should  be  made  to  the  leaflets  of  the  firms  supplying  the 
said  instruments.  By  the  use  of  the  paraboloid  condenser,  the  finer 
details  of  certain  stages  of  life-cycles,  such  as  the  formation  of 
granules  in  spirochaetes  and  treponemata,  can  be  observed  more 
readily  than  by  using  the  ordinary  substage  of  the  microscope. 
The  use  of  the  paraboloid  condenser  for  the  detection  of  small 
numbers  of  living  organisms  renders  it  of  value  for  rapid  diagnostic 
purposes. 

Stained  Material. 

Fuller  accounts  of  the  technique  of  fixed  and  stained  material 
will  be  found  in  Bolles  Lee  and  in  Langeron,  already  mentioned. 

Thin  Films. — For  the  examination  of  blood-inhabiting  Protozoa, 
it  is  necessary  to  make  first  thin  films  or  smears  of  blood.  There  are 
many  ways  of  doing  this,  and  opinions  differ  as  to  their  respective 
merits.  A  simple  method  is  to  take  a  straight  surgical  needle  about 
2  in.  long,  the  eye  of  which  has  been  removed,  and  a  clean  glass  slide. 
The  patieiit's  skin  is  pricked,  and  when  the  bead  of  blood  reaches 
the  size  of  a  small  pin's  head,  the  slide  is  applied  to  the  surface  of 
the  blood,  about  J  in.  from  the  far  (left-hand)  end  of  the  slide. 
The  shaft  of  the  needle  is  laid  across  the  drop  of  blood,  which 
spreads  between  the  slide  and  the  needle.  The  latter  is  drawn  evenly 
along  the  slide  towards  the  right.  The  film  is  dried  by  waving  it  in 
the  air.  The  film  should  possess  a  straight  edge  parallel  with  that  of 
the  slide  and  should  be  as  uniform  and  thin  as  possible.  Another 
glass  slide  may  be  used  as  a  spreader,  or  a  cover-slip  or  thin  glass 
rod  may  be  employed. 

Thick  Films. — These  are  of  service  in  detecting  malarial  paiasites 
or  trypanosomes,  especially  when  the  parasites  are  few.  The  method 
of  Ross,  or  a  modification  thereof,  has  been  much  used.  A  small  drop 
of  fresh  blood  is  spread  evenly  and  quickly  with  a  needle-point  over 
a  square  area  somewhat  less  than  that  of  an  ordinary  square  cover- 
glass.  The  blood  is  allowed  to  dry.  The  film  is  then  carefully 
dehaemoglobinized  in  water  in  which  there  is  a  trace  of  acetic  acid. 
The  dehaemoglobinizing  fluid  is  then  carefully  drained  off  and  the 

47 


748  THE   ANIMAL   PARASITES   OF   MAN 

film  again  dried.  It  is  fixed  in  absolute  alcohol  and  stained  with 
Romanowsky's  solution.  A  cubic  millimetre  of  blood  divided  into 
quarters  may  be  thus  dehaemoglobinized  and  stained.  The  parasites 
in  such  a  cubic  millimetre  of  blood  may  be  counted.  Such  a  proced- 
ure was  followed  by  R.  Ross  and  D.  Thomson/  in  determining  the 
periodic  variation  of  the  numbers  of  trypanosomes  in  the  blood  of 
a  patient,  as  mentioned  and  figured  on  pp.  78  and  79. 

For  cytologica  details  of  various  Protozoa,  thin  film  preparations 
on  cover-slips  or  slides  are  often  useful.  Cover-slip  preparations  are 
preferable,  unless  the  organisms  under  investigation  are  extremely 
scanty.  The  medium  containing  the  organisms,  such  as  blood, 
lymph,  intestinal  contents,  sputum,  scrapings  of  ulcers,  and  urine,  is 
spread  thinly,  either  alone  or  diluted  with  a  little  physiological  salt 
solution,  on  the  cover-slip.  Fixation  while  still  wet  is  necessary. 
Various  methods  are  employed. 

Fixatives. — A  useful  procedure  is  to  fix  the  wet  film  by  exposure 
to  4  per  cent,  osmic  acid  vapour  for  ten  to  thirty  seconds,  then  place 
in  absolute  alcohol  for  five  minutes  to  harden.  Grade  down  from 
absolute  alcohol  through  90  per  cent.,  70  per  cent.,  50  per  cent.,  and 
30  per  cent,  alcohols  to  water.  Stain  wet  with  a  suitable  stain  such 
as  haematoxylin,  and  gradually  dehydrate  by  grading  through  the 
necessary  strengths  of  alcohol,  clear  in  xylol  or  other  oily  clearing 
medium  and  mount  in  Canada  balsam. 

Other  fixatives  may  be  employed,  such  as  are  also  useful  for  fixing 
pieces  of  tissue  for  sectioning.  Films  or  smears  on  cover-slips  while 
still  wet  are  floated  on  the  surface  of  the  fixative  in  a  watch  glass. 
Some  good  fixatives  of  wide  application  are  : — 

Schaudinn's  Fluid. — This  consists  of  a  mixture  of 

Saturated  aqueous  solution  of  corrosive  sublimate  ...  ...       2  volumes 

Absolute  alcohol  ...  ...  ..,  ...  ...       i  volume 

Two  modifications  of  Schaudinn's  formula  may  be  found  useful. 
A  saturated  solution  of  corrosive  sublimate  in  physiological  salt 
solution  may  be  substituted  for  the  aqueous  one,  and  the  addition  of 
a  few  drops  of  glacial  acetic  acid  to  either  of  the  preceding  mixtures 
may  be  made. 

Some  workers  prefer  to  use  hot  fixatives,  raised  to  a  temperature 
of  about  50^"  C. 

Fixation  by  corrosive  sublimate  solutions  must  be  followed  by 
thorough  removal  of  the  mercury  salt  by  washing  repeatedly  in 
30  per  cent,  alcohol  or  with  iodine-alcohol. 

'  Froc.  Roy.  Soc.^  B,  Ixxxii,  p.  411. 


APPENDIX   ON    PROTOZOOLOGY  749 

Bouin's  Fluid,  or  modifications  thereof,  is  also  very  useful  for  wet 
fixation.     Bouin's  picro-formol  solution  consists  of  : — 

Saturated  aqueous  solution  of  picric  acid  ...         30  volumes 

Formalin,  40  per  cent.  ...  ...  ...         10       ,, 

Acetic  acid,  glacial  ...  ...  ...  2        ,, 

The  best-known  modification  is  one  due  to  Duboscq  and  Brasil, 
and  often  known  as  Bouin-Duboscq  Fluid.    Its  formula  is  as  follows  : — 

Alcohol,  80  per  cent.  ...  ...  ...  150  c.c. 

Formalin,  40  per  cent.  ...  ...  ...  60    ,, 

Acetic  acid,  glacial  ...  ..,  ...         15    ,, 

Picric  acid                ...  ...  ...  ...           i  grm. 

Thorough  washing  of  the  smear  or  cover-slip  preparation  with 
70  per  cent,  alcohol  until  the  yellow  colour  disappears  is  necessary 
to  remove  excess  of  fixative. 

Other  fixatives,  which  may  be  of  use,  more  especially  for  fixing 
small  pieces  of  tissue  for  sectioning,  are  the  solutions  of  Flemming 
(chromo-aceto-osmic  acids)  and  of  Zenker  (sublimate-bichromate- 
acetic,  with  sodium  sulphate). 

Regarding  the  time  of  fixation,  there  is  much  difference  of  opinion. 
Usually,  exposure  to  or  contact  with  the  fixative  for  five  minutes  is 
sufficient  in  the  case  of  films  or  smears.  Material  for  sections 
should  be  cut  into  small  cubic  pieces,  of  a  thickness  of  about 
5  mm.  (J  in.).  One  or  two  hours  should  be  sufficient  time  for  the 
fixation  of  such  pieces  of  tissue,  though  some,  as  Langeron,  prefer 
a  longer  time  of  fixation.  On  the  other  hand,  Gustav  Mann'  recom- 
mends a  short  fixation  period.  The  excess  of  fixative  should  be 
thoroughly  washed  out  of  the  tissue  in  the  manner  appropriate  to 
the  particular  fixative  used.  If  it  is  desired  to  keep  the  tissue  for  some 
time  before  sectioning  and  staining,  it  should  be  transferred  to  70  per 
cent,  alcohol. 

When  fluid  fixatives  are  employed,  large  quantities  of  the  fixing 
media  are  necessary.  The  volume  of  the  fixative  should  be  at  least 
ten  to  twenty  times  that  of  the  object,  and  the  latter  should  be  sus- 
pended in  the  middle  of  the  fixative.  The  tissue  should  be  fixed  as 
soon  as  possible  after  the  death  of  the  host. 

For  sectioning  tissue  parasitized  by  Protozoa,  embedding  in 
parafifin  is  generally  recommended.  Microtome  sections  should  not, 
if  possible,  exceed  5  //,  in  thickness.  Details  of  special  procedures 
must  be  sought  in  larger  works. 

Staining. — Here,  as  with  fixatives,  much  choice  is  presented.  The 
various  modifications  of  the  Romanowsky  stain  have  aided  greatly  in 

1  <<  Physiological  Histology,"  1902,  Clarendon  Press,  Oxford. 


750 


THE   ANIMAL   PARASITES   OF   MAN 


the  detection  of  various  Protozoa  parasitic  in  the  blood.  Such 
stains,  however,  leave  something  to  be  desired  in  the  revealing  of  finer 
cytological  details.  Other  stains,  more  especially  the  haematoxylins, 
must  be  employed  for  cytological  purposes. 

FormulcC  of  some  of  the  principal  Romanowsky  and  ha^matoxylin 
stains  may  now  be  given. 

The  underlying  principle  of  the  Romanowsky  Stain  is  the  reaction 
between  alkaline  methylene  blue  and  eosm,  forming  the  so-called 
eosinate  of  methylene  blue  which  stains  chromatin  purplish-red. 
A  solution  of  medicinal  methylene  blue  after  having  been  subjected 
to  the  action  of  an  alkali,  such  as  sodium  carbonate,  becomes  partly 
converted  into  certain  derivatives,  the  chief  of  which  are  methylene 
azure  and  methylene  violet.  These  substances  are  also  present  in 
matured   polychrome   methylene    blue. 

The  formula  of  a  slightly  modified  Romanoivsky  Stain  which  gives 
excellent  results  is  given  below  : — 

Two  stock  solutions  are  required — 

Solution  A. — Methylene  blue,  pure  medicinal  ...  ...  i*o  grm. 

Sodium  carbonate    ...  ...  ...  ...         0*5     ,, 

W^ater         ...  ...  ...  ...  ...     loo-o  c.c. 

Keep  in  a  warm  incubator  for  two  or  three  days,  until  the  solution  is  distinctly  purple  in 
colour.     It  improves  with  age. 

Solution  B. — Eosin,  water  soluble,  extra  B. A.  ...  ...         10  grm. 

Water         ...  ...  ...  ...  ...   1,000 'O  c.c. 

This  solution  must  be  kept  in  the  dark,  in  dark-tinted  (amber-coloured)  bottles,  as  unfortu- 
nately it  is  decolorized  by  light. 

Before  use  each  stock  solution  must  be  diluted.  Thus,  make  up 
5  c.c.  of  each  stock  solution  to  100  c.c.  by  adding  distilled  water. 
For  staining,  i  volume  of  solution  A  is  added  to  2  or  3  volumes  of 
solution  B.  Mix  thoroughly  by  shaking,  pour  the  mixture  over  the 
film,  previously  fixed  in  absolute  alcohol,  and  stain  for  ten  to  fifteen 
minutes.  Wash  carefully  in  running  w^ater,  then  dry.  The  cytoplasm 
of  a  protozoan  parasite  will  be  stained  blue,  the  chromatin  purplish- 
red  and  vacuoles  or  very  tenuous  protoplasm  wall  remain  colourless. 

The  exact  proportions  of  solutions  A  and  B,  which  must  be 
mixed  together,  should  be  determined  by  experiment.  Freshly  mixed 
stain  must  be  used  on  each  occasion. 

Leishman's  Stain  is  the  precipitate  resulting  from  the  interaction 
of  alkaline  methylene  blue  and  eosin.  The  washed  and  dried 
precipitate  is  collected  and  dissolved  in  pure  methyl  alcohol,  which 
acts  as  a  fixative  ;  0*015  grm.  of  Leishman  powder  may  be  dissolved 
in  10  c.c.  of  methyl  alcohol  for  staniing  films.  The  film  is  covered 
with  the  solution  for  one  minute,  twice  the  volume  of  water  is  then 
added  and  mixed  with  the  stain   on  the  slide.     The  staining  is  then 


appendix:  on  protozoology  751 

continued  for  five  to  ten  minutes,  and  the  film  is  finally  washed  with 
water. 

Gienisii's  Stain.  —  This  should  be  procured  ready  made.  Azure  II 
is  a  mixture  of  methylene  azure  and  methylene  blue.  (Methylene 
azure  is  sometimes  known  as  Giemsa's  Azure  I.)  The  formula  given 
by  Giemsa  himself  in  19 12  is  : — 

Azure  Il-eosin           ...  ...  ...  ...         3*0  grm. 

Azure  II     ...              ...  ...  ...  ...         0'8    ,, 

Glycerine,  pure         ...  ...  ...  ...  125*0    ,, 

Methyl  alcohol,  pure  ...  ...  ...  375"0    ,, 

The  film  is  first  fixed  in  absolute  alcohol.  The  proportion  of 
stain  usually  used  is  one  drop  of  stain  to  i  c.c.  of  water.  Stain  for 
about  ten  minutes  and  then  wash  in  water. 

The  details  of  the  application  of  the  Giemsa  stain  to  films  fixed 
wet  and  to  sections  must  be  sought  in  larger  works  on  technique. 
These  works  should  also  be  consulted  for  information  regarding 
the  use  of  Pappenheim's  Panchrome  mixture. 

There  are  numerous  formulae  of  stains  containing  ripened  Hama- 
toxylin  or  its  essential  principle,  Hceniatein.  A  mordant  is  necessary, 
one  of  the  alums  being  usually  employed.  The  mordant  may  be 
included  as  an  ingredient  in  the  staining  mixture,  or  it  may  be  used 
separately  as  in  the  case  of  the  so-called  iron-haematoxylins,  wlierein 
ferric  ammonium  alum  is  used  separately  and  is  followed  by  staining 
with  haematoxylin  or  haematein.  A  few  of  these  stains  of  general 
application  may  now  be  mentioned. 

Delafield's  {or  Grenachers)  Hcematoxylin, 

Hgematoxylin  crystals              ...  ...  ...         4  gi^m. 

Absolute  alcohol       ...             ...  ...  ...  25  c.c. 

Saturated  aqueous  solution  of  ammonia-alum  ...  400    ,, 

Mix  these  ingredients,  and  leave  exposed  to  light  and  air  for  three  to  four  days.    Filter 
and  add — 

Glycerine   ...             ...             ...  ...  ...  100  c.c. 

Methyl  alcohol          ...             ...  ...  ...  ico   ,, 

Allow  the  mixture  to  stand  until  the  colour  is  sufficiently  deep, 
then  filter  and  place  in  a  stoppered  botde.  The  solution  should  be 
allowed  to  ripen  for  at  least  two  months  before  use.  Dilute  aqueous 
solutions  of  the  stain  are  of  service  for  films  and  for  sections.  A 
trace  of  acetic  acid  may  be  added  at  the  moment  of  use,  for  sharp 
differentiation. 

Ehrlich's  acid  haematoxylin,  Mayer's  ha^malum,  and  Mayer's 
glych^malum  are  also  useful.  Their  formulae  will  be  found  in  larger 
works. 


75^ 


THE   ANIMAL   PARASITES   OF   MAN 


The  chief  Iron-H cematoxylin  Stain  is  that  devised  by  Heidenhain. 
Unfortunately  the  procedure  involved  is  a  long  one,  and  various 
modifications  have  been  made  to  obviate  this  disadvantage.  Hsema- 
tein  maybe  used  instead  of  ripened  haematoxylin. 

One  efficacious  modification  of  Heidenhain's  stain  is  that  of 
Rosenbusch.  The  smear  or  tissue,  after  fixation,  must  be  graded 
downwards  through  the  alcohols  to  water.  Mordant  for  one  and  a 
half  hours  in  a  3J-  per  cent,  aqueous  solution  of  ferric  ammonium 
sulphate.  Stain  for  about  three  minutes  in  i  per  cent,  solution  of 
ripe  haematoxylin  or  haematein  in  absolute  or  96  per  cent,  alcohol, 
to  which  a  drop  of  saturated  aqueous  sokition  of  lithium  carbonate, 
sufficient  to  produce  a  wine-red  colour,  has  been  added.  Differentiate 
under  the  microscope  with  a  very  dilute  solution  of  the  ferric 
ammonium  sulphate.  Wash,  gradually  dehydrate,  clear  and  mount 
in  balsam.  It  must  be  remarked  that  iron-haematoxylin  is  a  regressive 
stain,  hence  great  care  must  be  exercised  in  differentiating  with  the 
iron  alum. 

Gentian  Violet. — A  i  per  cent,  alcoholic  solution  of  gentian  violet, 
or  of  methyl  violet,  or  of  crystal  violet,  will  be  found  useful  for  stain- 
ing spirochaetes. 

Methyl  Green. — This  substance  is  considered  to  be  a  chromatin 
stain,  for  either  fresh  or  perhaps  recently  fixed  tissues.  A  concen- 
trated aqueous  solution  contains  about  i  per  cent,  of  the  stain.  This 
should  be  added  to  a  i  per  cent,  solution  of  acetic  acid.  It  may 
be  used  for  demonstrating  the  nuclei  of  ciliates. 

In  conclusion  it  is  essential  to  remember  that  the  actual  magnifica- 
tion of  figures  of  Protozoa  should  be  given,  and  not  merely  the 
combination  of  objective  and  ocular  that  has  been  used,  for  unless 
the  tube-length  and  distance  of  the  drawing  board  from  the  ocular  be 
also  given,  it  is  not  possible  to  compute  the  magnification  from  such 
information.  Drawings  should  always  be  made  with  the  aid  of 
a  camera  lucida,  drawing  prism  or  other  form  of  projection  apparatus. 


APPENDIX   ON   TREMATODA   AND   NEMATODA  753 


APPENDIX  ON  TREMATODA  AND 
NEMATODA. 

BY 

J.  W.  W.  STEPHENS,  M.D.,  B.C.,  D.P.H. 
TREMATODA. 

Artyfechinostomum  sufrartyfex,  Clayton  Lane,  19 15. — Leiper 
thinks  this  may  be  the  same  as  Echhiostoma  jualayamnn,  Leiper, 
1 91 1,  which  species  Odhner  assigns  to  the  genus  Euparyphium. 

Metagonimus  (Yokogawa)  yokogawai  occurs  in  dogs  in  Shanghai. 
Encysted  cercariae  probably  in  the  perch. 

Opisthorchis  sp. — Skin  covered  with  spines.  Gut  forks  almost 
reach  end  of  body.  (Esophagus  two  to  three  times  length  of 
pharynx.  Ovary  multilobed.  Ovary  and  testes  in  posterior  fourth 
of  body.  Vitellaria  end  opposite  the  ovary.  Distinguished  from 
0.  felineus  by  presence  of  spines  and  lobed  ovary ;  from  0.  pseudo- 
felineus  and  0.  noverca  by  the  lobed  ovary,  and  by  the  fact  that  the 
yolk  glands  do  not  extend  as  far  as  the  anterior  testis.  It  agrees 
with  Poirier's  description  of  0.  viverrini  in  the  Indian  civet  cat,  but 
whether  this  species  has  spines  on  the  cuticle  is  not  known. 

Habitat. — Man  in  Chiengmai  (Malay  States).  Fifteen  per  cent, 
of  prisoners  in  the  jail  showed  the  ova  of  this  species  in  their  faeces. 

Schistosome  cercariae. 

Schistosome  cercariae  belong  to  the  furcocercous  division  of  the 
Distomata  cercariae. 

Distomata  cercariae. 

Body  without  a  floating  membrane.  Tail  absent,  or  if  present 
not  cleft  to  the  base.  Mouth  anterior,  gut  forked.  Oral  sucker 
present.    Ventral  sucker  near  middle  of  body.     Eyes  generally  absent. 

Group     Fercocercous  cercariae, 

Cercariae  single  (not  in  colonies).     Tail  forked  at  its  end. 

Family.     Schistosomidae. 

Pharynx  absent. 


754  THE   ANIMAL    PARASITES   OF   MAN 

Cercaria  bilharzia,  Leiper,  1915. 

Pigment  spots  (eyes)  anterior  to  ventral  sucker  absent,  cuticular 
keel  on  forks  of  tail  absent. 

In  Bullinus  sp.  and  Planorbis  hoissyi  in  Egypt,  (?)  in  Physopsis 
africanuy  South  Africa.     Adult  form,  Schistosoma  hcematohium. 

Cercaria  bilharziella,  Leiper,  1915. 

Cuticular  keel  on  tail  forks  present.  Pigment  spots  (eyes)  in  front 
of  ventral  sucker  present. 

In  Planorbis  hoissyi  and  P.  mareoticus,  and  in  Melanin  sp.  Adult 
form(?). 

For  characters  of  numerous  other  cercariae  which  occur  in  fresh 
water  molluscs  see  "  Die  Susswasserfauna  Deutschlands,"  Max  Liihe, 
H.  17  (Gustav  Fischer,  Jena,  1909). 

The  characters  of  Cercaria  japonica  of  S.  japonicuni  in  the  mollusc 
Katayama  nosophora  and  of  C.  mansoni  have  still  to  be  defined. 

Schistosoma  mansoni,  Sambon,  1907. 

The  evidence  appears  to  be  strong  that  terminal-spined  eggs  are 
not  found  in  the  West  Indies,  and  that  therefore  the  lateral-spined 
eggs  found  in  faeces  there  belong  probably  to  S.  mansoni.  If  this 
be  true,  then  the  egg  described  by  Stephens  and  Christophers 
in  man  in  India  probably  also  belongs  to  another  species  of 
Schistosome. 


NEMATODA. 

Ancylostomiasis.  —  Treatment:  (i)  Oleum  chenopodii  (U.S. P.), 
dose  iri  X  to  irt  xv  on  a  lump  of  sugar,  three  doses  at  two-hourly 
intervals,  preceded  and  followed  by  a  purge.  It  is  cheap,  not 
unpleasant  to  take,  and  non-toxic.  Effective  also  against  Ascaris 
lumbricoides. 

(2)  Milk  of  the  higueron  Ficns  laurifolia.  A  spoonful  in  milk, 
three  times  daily  for  three  days  followed  by  a  purge.  Described  as  a 
harmless  but  very  sucessfu    form  of  treatment. 

Ground-itch. — Completely  cured  in  a  few  days  by  a  3  per  cent, 
solution  of  salicylic  acid  in  ethyl  alcohol.  Apply  for  five  minutes 
twice  daily. 

Ascaris  lumbricoides  can  be  kept  alive  for  twelve  days  in 
Kronecker's  solution  ;  NaHO  0069  grammes,  normal  saline  1,000  c.c. 

Eggs  are  laid  and  develop  in  about  a  fortnight  at  ordinary  room 
temperature.     At  70"^  C.  they  are  readily  killed. 


APPENDIX   ON   TREMATODA   AND   NEMATODA  755 

Filariasis. —  Dutcher  and  Whitmarsh  have  cultivated  from  the 
blood  and  from  the  exudation  fluids  of  cases  of  filariasis  (elephanti- 
asis, lymphangitis,  etc.),  in  about  sixteen  cases,  a  bacillus  resembling 
B.  subtil  is.  Controls  were  negative.  They  propose  the  name  Bacillus 
lymphangiticus  for  this  organism,  and  they  believe  it  to  be  the  cause 
of  the  diseases  grouped  under  the  designation  "  filariasis." 

Oncocerca  volvulus. — Unsheathed  embryos  (indistinguishable  from 
those  taken  from  the  uterus  of  this  worm)  have  been  found  in  lymph- 
atic glands  and  in  the  blood  (if  considerable  pressure  is  used  so  as 
to  squeeze  out  lymph  at  the  time  of  taking  the  linger  blood,  otherwise 
none  occurs  in  the  specimens).  The  measurements  in  dried  films 
are  :  Nerve  ring  237  per  cent,  of  length  ;  Gi  cell  69-6  per  cent.  ; 
end  of  last  tail  cell  96-3  per  cent;  total  length  274-3  /n. 

Strongyloides  stercoralis. — Pathology  :  They  occur  in  the  wall 
of  the  intestine  and  may  be  associated  with  ulceration.  They  also 
occur  in  lymphatics  and  blood-vessels. 


756  THE   ANIMAL   PARASITES   OF   MAN 


BIBLIOGRAPHY. 

[In  the  following  pages  the  letters  C.  f.  B.,  P.  u.  Inf.  are  used  to  indicate 
the  Centralhlatt  fur  Bakteriologie_,  Pathologie  und  Infektions-Krankheiten.'] 

(A)  PROTOZOA  (pp.  25  to  210,  617  to  637,  and  y^z  to  742). 
[This  list  affiles  to  the  earlier  literature  only.     More  recent  references  are 
given  as  footnotes  in  the  text.'] 

(a)  General. 
BtJTSCHLl,  O.     Protozoa  in  Bronn's  Klass.  u.  Ordn.  d.  Tierreichs,  Leipz.,  1880- 

1889. 
Calkins,  G.  N.     The  Protozoa,  Columbia  Univ.  Biol.  Ser.,  vi.  New  York,  iqoi. 
Delage,  Y.,  and  E.  Herouard.      Traite  de  Zool.   Concr.,   i,  La  cellule  et  le& 

protozoaires,  Paris,  i8q6. 
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A  Treatise  on  Zoology,  edited  by  E.  Ray  Lankester,  London,  1903,  i,  2. 
Lang,   A.     Lehrb.    d.   vergl.   Anatomic   d.    wirbellos.    Tiere,   2.   Aufl.,  2.  Lief,. 
Protozoa,  Jena,  1901. 

(b)  Pathogenic  Protozoa  in  General. 
Doflein,   F.     Die   Protozoen   als  Parasiten  u.   Krankheitserreger,  Jena,    1901 ;. 

Lehrbuch  der  Protozoenkunde,  1912. 
Doflein,   F.,   and   S.   v.    Prowazek.      Die  pathog.    Protoz.    mit  Ausnahme  d. 

Hamospor.,  in  Handb.  d.  path.  Mikroorganism.  ;  issued  by  W,  Kolle  and 

A.  Wassermann,  11.  and  12.  Lief,  Jena,  1903. 
Kastner,  p.     Die  tierpathogenen  Protozoen,  Berlin,  1906. 
KISSKALT,    K.,  and   M.    Hartmann.      Praktikum  der  Bakteriologie  und  Proto- 

zoologie,  Jena,  1907. 
LtJHE,   M.     Die  im   Blute  schmarotzenden   Protozoen  und   ihre  nachsten   Ver- 

wandten,  in  Handb.  d.  Tropenkrankh.,  issued  by  C.  Mense,  Leipz.,  1906,, 

iii. 
Pfeiffer,  L.     Die  Protozoen  als  Krankheitserreger,  2.  Aufl.,  Jena,  1891 ;  Supple- 
ment, Jena,  1895. 
Roos,    E.     Die  im   menschl.    Darm    vork.    Protozoen   u.    ihre   Bedeutg.,    Med. 

Klinik,  1905,  i,  p.  1328. 
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Haustiere,  Leipz.,  1898. 
SlEVERS,  R.     Zur  Kenntn.  d.  Verbreitg.  d.  Darmparas.  d.  Mensch,  Helsingfors, 

1905,  Festschrift  f.  Palmen. 
Ward,  H.  B.     Protozoa,  Wood's  Ref.  Handbook  of  the  Med  Sci.,  1904,  viii. 

Class  I — Sarcodina  (pp.  29  to  50). 

Order.     Amoebina  (p.  29). 

Entamoeha  coli;  Entamceba  histolytica   (pp.  32  to  41,  618  to  620,  and  y2>3). 

Blanchard,  R.     Traite  de  Zool.  medic,  1885,  i,  Paris,  p.  15. 
—  Maladies  paras.,  1895,  p.  658. 

Bowmann,  M.  H.     Dysentery  in  the  Philippines,  Journ.  Trop.  Med.,   1901,  iv,. 
p.  4^0. 


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■^58  THE    ANIMAL    PARASITES    OF    MAN 

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y6o  THE   ANIMAL   PARASITES   OF   MAN 

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[An  almost  complete  collection  of  the  literature  relating  to  Cestodes  up  to 
1895   is  to    be    found   in    Braun's    monograph   on    the  tapeworms    in    Bronn's 
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79C 


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836 


THE   ANIMAL   PARASITES   OF   MAN 


INDEX 


Abdomen,  malignant  growth,  in,  with 
ascites,  Leydenia  gemmipara  associated 
with,  49,  50 

Abdominal  and  j^elvic  organs,  blood-supply 
of,  as  illustrating  distribution  of 
Schistosoma  hcematobium  in  body,  272 

—  operation    wound,    escape    of    Ascaridea 

from,  654,   655 
Abscess  cavities,  larvae  of  Sarcophaga  wohl- 
fahrti  from,   723 

—  in  filariasis,  401 

Abscesses,  cutaneous,  due  to  Lagocheilas- 
caris  minor,  467 

Abyssinians,  infection  with  Tcenia  saginata, 
340 

Acanthiadoe,  characters,  534 

Acanthia  lectularia,  see  Cimex  lectularius 

Acanthobothrium  coronatum,  excretory  ves- 
sels, 292 

Acanthocephala,   475 

—  development  of,    17 

—  isolated  position  of,  20 

— -  life  spent  in  intermediate  and  final  host, 

18 
— ■  loss  of  digestive   system   in,  3 

—  mori:)hology,  475 
Acanthocheilonema,  morphology,  414 

—  perstans,   geographical    distribution,   416 
in  South  America,  416 

morphology,  414 

topographical  distribution,  416 

Acarina,  characters  and  morphology,  484 

—  hosts,  habitat  and  food  of  (and  footnote), 

484 
Acartomyia,   characters,  564 
Acarus  dysentericB,  512 

—  hordei,  cereal  mite,  489 
Accessory  sinuses,  nasal,  larvae  in,  717 
Acephalina  often  "  coelomic  "  parasites,  134 
Acid  alcohol  in  differentiation  of  flukes,  471 
Acinetaria,    198 

Acne,  lesions  resembling,  set  up  by  Sparga- 

nuni  proliferum,  318 
Acoleince,  vagina  atrophied  in,  297 
Actinomyxidia,    129,    187 
Adams,    skin    disease    caused    by    larvae    of 

Dermatobia  noxialis,  725 
Addario,    Filaria    (?)    conjunctivce    in    man, 

405 
Adelea,  141 
Adeleidea,    141,    742 
Adie,  Mrs.,  life-cycle  of  Hcetnoproteus  (Hal- 

teridiiim)   columboe,   152 
iEdeomyia,  characters,  565 


I   ^deomyina,  characters,  564 

I    iEdes,  characters,  564 

I    Africa,   larvae   of  Muscidce   causing   myiasis 

i  in  man  in   (footnote),  590 

i   —  Sduth,  farm  stock  in,  attacked  by  Hya- 

I  lom,ma  cegyptium,  501,  502 

'   —  West,  French,  cause  of  myiasis  in,  614 

I geographical    distribution    of    Oncho- 

I  cerca  volvulus  in,  419 

I   Agamo filaria,  406 

—  georgiana,  habitat,  406 
morphology,   406 

,   —  labialis,  morphology,  407 

—  oculi  humani,  405,  406 

—  palpebralis,   405,  406 
j    Agamomermis,   470 

j   —  restiformis ,  morphology,  470 

Agglutinating  haemolytic  action  of  serum  of 
j  ancylostome    patients,   648 

[   Ahmed  Emin,  small  variety  of  Plasmodium 
I  vivax,   166 

Air-passages,  invasion  by  larvae  of  screw- 
worm  fly  (Chrysomyia  macellaria),  587 

by  Metastrongyliis  apri,  433 

—  upper,  Ascarides   in,   690 

danger  of,  691 

how  introduced,  690 

leeches   in,   699,    700 

Aitken,  Porocephalus  constrictiis ,  526 

Akamushi,  see  Kedani  mite 

Akaneesch,  see  Kedani  mite 

Alcohol,  application  in  nasal  myiasis,   719 

Aleppo  button,  see  Oriental  sore 

Aleurobius  (Tyroglyphus)  farince,  charac- 
ters of,  511 

Alexeieff,  Chilomastix  (Tetramitus)  mesnili, 
735 

—  on    genus    Giardia,    736 

—  views   on   trichomonad   cysts,   56 
Alimentary  canal  of  Hirudinea,  480 

—  tract,  spirochaetes  in,  741 
Allessandri,    haemolytic    action    of    ancylo- 

stomes,   647 

Allocreadiuin  isoporum,  excretory  appara- 
tus, 218 

Alum  solution  in  nasal  myiasis,  719 

Amaurosis  following  male  fern  poisoning, 
671 

Amblyomma  americana,  geographical  distri- 
bution, 501 

objects  of  attack,  501 

suppuration  resulting  from  punctures 

of,  501 

—  cayennense,   characters,   500 
geographical  distribution,  501 

—  —  ill-effects  from  attacks  of,  501 


INDEX 


•37 


Amhlyomma    cayennense,    synonyms,    500 

—  characters  of,  497 

—  hebroeum    (bont    tick)    carrier    of    heart- 

water    fever   in    sheep,   493 
life-cycle   of,   495 

—  maculatum,  501 

America,    North,    amount   of   prevalence    of 
trichinosis   in,  428 

—  South,      geographical      distribution      of 

Acanthocheilonema  perstans    in,    416 
Amicis,  de,  infection  with  Demodex  follicu- 

lorum,  708 
Amoeba   buccalis,  44,  734 

—  coli,   31 

—  dentalis,  44,  734 

—  diaphana,  31 

—  dysenterioB,    31 

—  fluid  a,    46 

—  gingivalis,  44,  734 

—  lobosa,  var.  guttula,  var.  oblonga,  31 

—  miurai,   46 
characters,  46 

—  reticularis ,  31 

—  spinosa,  31 

—  urogenitalis,  45 

—  I'ermicularis,  31 

Amoebae,  bodies  resembling,  found  in  serous 
fluid,  46 

—  cerebral    abscess  set  up  by,   35 

—  cultural,    42,    618 

—  culture  media  for,   742,  743 

—  experimental    injection    of,    in    cats    pro- 

ducing dysentery,  35 
—  producing  enteritis,   36,   37 

—  experiments    with,    618 

—  found  in   faeces,  47,  48 
in   lung,   45 

in   urine,   45,   46 

—  intestinal,  association  with  colitis,  30 
causal  agent  in   production  of  dysen- 
tery, 30 

—  experiments       made        to 

prove,    30 

culture  medium   for,   743 

— -  —  discovery  in  case  of  dysentery,  29,  30 

—  in    stools,    30 

encysted,  31 

human,  29 

discovery    of,    29 

ingestion  of  red  blood  corpuscles  by, 

35,  39,  42 

—  invading  bladder,  46 

—  liver   abscess  set  up  by,  35 

—  penetration     of     intestinal     blood-vessels 

by,  36 

—  phagedsenic,   733 

—  pulmonary  abscess  set  up  by,  35 

—  testaceous     (Monothalamia),     characters 

of,  47 
Amoebic    dysentery,    618 
Amoebina,  characters  and  habitat,  27 
Amoeboid  germs  in  pseudonavicellae,  130 
Amoebosporidia,    130 
Amoebulae,  34,   49.   183,   185 

—  formation  of,  34 
Amphimerus,  257 


Amphimerus    noverca,    habitat,    258 

morphology,    257,   258 

synonyms,  258 

Amphistomata,  morphology,  230 
Anaemia,    case    of,    effect    of    expulsion    of 
Taenia  solium  on,  648 

—  in  ancylostomiasis,  treatment,  687 

—  of  dourine,  97 

—  period  of,  in  liver-fluke  disease  in  sheep,. 

240 

—  pernicious,  symptoms  disappearing  after 

expulsion  of  AscaridcB,  049 

—  proteid  metabolism  in,  645 

—  splenic,    infantile,   see   Kala-azar,   infan- 

tile 

—  see  also  Ancylostome  anosmia 

—  see  also  Bothriocephalus  anaemia 

—  see  also  Trichocephalus  anosmia 

Anal  canal,  means  of  access  of  Schistosoma 

hcematobium  to,   272 
Anaplasma,    180 

—  marginale,    180,   611 

cause  of  gall-sickness  in  cattle,  180 

Ancylostoma  braziliense,  456 

—  caninum,  456 

—  ceylanicum,  habitat,  456 

—  —  morphology,  456 

—  characters,  445 

—  duodenale  and  Strongyloides  stercoralis y 

larvae  of,  differences  hjetween,  451 

buccal  capsule,  445 

bursa,   448,   449,   450 

cephalic    glands,    447 

cervical  papillae,  447 

—  —  development,  451 

diagrammatic  representation  of  male 

and  female,  446 

embryos,    451 

excretory  system  and  cervical  glands, 

447 

diagrammatic  representa- 
tion,  448 

food,  450 

genital   cone,   450 

geographical  distribution,  450 

habitat,  in  man  only,  450 

infection   by,   682 

modes  of,  683 

must  be  on  large  scale  to  produce 

illness,  682 

see  also  Ancylostomiasis 

invading  frontal  sinus,  683 

—  —  larvae   of,   bionomics   of  development, 

453,  454 

infection  by  skin,  454,  455 

in  dogs,  455 

infective  stage,  454 

method  of  cultivation,  455 

mode  of  entry  into  body,  454 

morphology,  451,  452,  453 

stages,   451,    452 

thigmotropism  in  mature  stage,  454 

lateral  lines,  448 

morphology,  445 

number  of  females  present  in  intes- 
tine, mode  of  reckoning  (footnote),  454 


838 


THE    ANIMAL    PARASITES   OF    MAN 


Ancylostoma  duodenale,  oesophageal  glands, 
447 

organs  of  Necator  americanus  com- 
pared with  those  of,  458 

ova   of,   451 

ovaries,   449 

spicules,   450 

testis,  449 

—  —  ventral  teeth,  446,  447 

—  larvae  of,  cultivation,  474 

—  malayanicum,   456 

■ —  pluridentatum,  456 

Ancylostome   anaemia,   etiology,   647,   648 

experimental,  646 

retinal   haemorrhages   in,   646 

toxic  hypothesis,  646,   647 

treatment,   687 

Ancylostomece,  445 

Ancylostomes,  expulsion  of,  drugs  for,  685, 
686 

—  toxic  action  on  hosts,  647 
Ancylostomiasis,    agglutinating    haemolytic 

action  of  serum  of  patients,  648 

—  eosinophilia  in,   647 

—  morbid  anatomy,  459 

—  prophylaxis   against,   684 
in  miners,  684 

—  proteid   destruction    in,    647 

—  symptoms,  683 

set    up    by    invasion    by    Hcemonchus 

contortus  mistaken   for  those  of,  438 

—  treatment,  754 
Ancylostominoe,  438 

Andrews,  Oxyuris  in  appendix,  655 

Angiostomidas,  374,  379 

Angiostomum      nigrovenosum,      heterogony 

in,  381 

male  of  rhabditic  form,  370 

mode  of  generation,  372 

Angola,   highlands  of,   uncertain   species  of 

Ascaris  occurring  in,  465 
Anguillula,  379 

—  aceti   found    in   vinegar,   379 
morphology,   379 

occurrence  in   urine,   379 

—  intestinalis      and      A.      stercoralia ,      see 

Strongyloides   stercoralis 

—  mucronata,  377 
AnguillulidcB,  377 

—  hosts  of,  374 

—  morphology,  374 
Anguillulina,  379 

- —  putrefaciens  living  in  oniors,  379 

synonyms,   379 

Animal  matter,  decomposing,  Tyroglrphidce 
in,   511 

Animals,  mites  living  endoparasitically  in, 
491 

Ankylorhynchus,  characters,  563 

Annaratone,  pseudomeningitis  due  to 
Ascajidce    infection,    649 

Anopheles  and  Culex,  larvae  of,  position  in 
water   compared,   554 

ova  of,  method  of  depositing  com- 
pared,  554 

points   of  difference  between,   551 


Anopheles  hijurcatiis,  ova  of,  localities 
selected  for  deposition,  553 

—  characters,  561,  566 

—  claviger,  mouth  parts,  550 

—  development  of  human  malarial  parasite 

only  takes  place  in,  158.   159 

—  entire     genus     capable     of    transmitting 

malaria  to  man,   552 

—  head  of  male  and  female,  549,  556 

—  only    genus     of    mosquito    transmitting 

malaria,    158 

—  ova  of,   557,   558 

—  maculipennis,  552 

breeding  places  of,  557 

Crithidia  inhabiting,   104 

intestine  of,  stages  of  development  of 

pernicious  or  malignant  tertian  para- 
sites in,  162 

larva  of,  553 

ova  of,  localities  selected  for  deposi- 
tion, 553 

pupa  of,  554 

sporulation  stages  of  malarial  para- 
sites from,  163 

stomach  of,  oocysts  and  ookinetes  of 

malignant  tertian  parasite  in,   162,  163 

transverse  section  through  proboscis, 

550 

Anophelines,  genera  of  (footnotes),  562,  563 

—  larvae    of,    destruction    in    prevention    of 

malaria,  636 

—  number  of  species,  552 

—  ova  of,  best  known,  559 
Anoplura,  532 

—  see  also  Pediculidce 

Antelope  tolerant  to  trypanosomes,  69 

—  Trypanosoma  gambiense  in,  76 

—  T.  rhodesiense  in,  69,  70 

Anterior  station  in   Glossina  of  a  trypano- 

some,    101 
Anthomyia  desjardensii,  cause  of  intestinal 

and  cutaneous  myiasis,  585 

—  pluvialis,  larvae  of,  584 

—  scalaris,      maggots      of,      passed      from 

urethra,  728 
AnthomyidcB,   flies   belonging   to,   attacking 

man,  611 
Anthrax  transmitted  by  Stomoxys,  610 
Antimony,  use  in  sleeping  sickness,  623 

—  and   atoxyl,   combined,   in   sleeping  sick- 

ness, 622 
Anus,    exit   of    Oxyuris    vermicularis   from, 
467 

—  prolapse    of,    set    up    by    migrations    of 

Oxyuris  vermicularis,  695 
Aphaniptera  (fleas),  characters,  543 

—  see  also  Fleas 

Aphides    (or  plant  lice)   (footnote),  532 

said  to  have  been  passed  in  human 

urine  (footnote),  532 
Aphiochceta  ferruginea,  582,  583 

—  —  characters,  583 

geographical  distribution,  583 

larvae  (maggots)  of,  583 

Aponomma,  characters  of,  497 

—  hosts  of,  497 


INDEX 


839 


Apoplexy  cause  of  death  in  first  period  of 
liver-fluke  disease  in  sheep,  240 

Appendicitis,  association  of  Oxyuris  vermi- 
cularis  with,  467 

—  in  relation  to  intestinal  parasites,  views 

of  authors  regarding,  652,  653,  654,  655 

—  relationship  of  Oxyuridce  to,  698 
Appendicostomy    in    gangrenous    dysentery, 

619 
Appendix   vermiformis,    ascaris   in,    causing 
intestinal    obstruction,   654 

—  —  bilharziasis  of,  642 

intestinal  parasites  invading,  authors 

recording  cases  of,  652 

Oxyuris  in,  654,  655 

perforation  by  Ascaris,  raritv  of,  655, 

656 

trichocephali   in,   655 

Aptera,   531 

Apterygota,  531 

Arachnoidea,  characters,  483,  484 

—  orders  of  (footnote),  484 

- —  relation  to  Linguatulidoe,  19 
Aradidce,  characters,  541 
Aragao,    on    Chlamydozoa,    209 

—  on    Hajmoproteus,    152 

—  on    leucocytogregarines    in    birds,    155 
Archigetes,     attains     maturity     in     lower 

animals,  21 

—  stage  of  sexual  maturity,  305 
Argantince  and  IxodincB,  distinguishing  fea- 
tures between,  505 

—  characters  of,  496 
Argas  brumpti,  507 

—  chinche,   508 

—  persicus,   506 

appearance  of  SpirochoBta  gallinarum 

in  hsemoccelic  fluid  of,  119 

bite  of,  serious  effects,  507 

blood-sucking,   507 

granules  in  digestive  tract  of,  507 

hosts  of,  507 

transmission  of  Spirochoeta  gallina- 
rum by,   119 

—  reflexiis,   bite   of,   symptoms   set   up   by, 

506 
blood-sucking  habits  of,   506 

—  —  geographical  distribution,  506 
habitat,   506 

vitality   of,   506 

—  species  of,  Neumann's  table,  505 
Arilus  carinatus,  542 

Avion  sp.,  scolex  of  cysticercoid  from,  with 
excretory   vessels   outlined,  292 

Armadillo,  possible  reservoir  of  Trypano- 
soma  cnizi,   87 

Arribalzagia,  characters,  562,  568 

Arsenic  and  glycero-phosphates  in  bronchial 
spirochaetosis,   633 

—  in  sleeping  sickness,  622,  623 

—  in   treatment  of  nagana,  94 
Arsenious    acid   in    bronchial   spirochaetosis, 

633 
Arseno-phenyl-glycin    in    sleeping    sickness, 
623 


Arseno-phenyl-glycin,  resistance  of  Try- 
panosoma  lewisi   to,    how   lost,   93 

Arslan,  experimental  ancylostome  anaemia, 
646 

Artault,  Entamoeba  pulmonalis ,  45 

Arthropoda,  483 

—  natural  flagellates  of,  104 

—  pebrine  bodies  or  Microsporidia  in,   184 

—  segmented  structure  of,  483 

—  skin  of,  how  hardened  (footnote),  483 
Artyfechinostomum,  morphology,  269 

—  sufrartyfex,   753 

morphology,   269 

Ascariasis,    diagnosis,    692 

Epstein's  method,  692 

AscaridoB,   375,  461 

—  chemically  toxic  effects  of,  650 

—  epidermal    cells,   isolated,   of    (footnote),. 

361 

—  expulsion  of,  favourable  effects  of,  649^ 

650 

—  infection   by,   causing  pseudomeningitis^ 

649,  650 
Ascarides  causing  constipation,   657 
intestinal  obstruction,  657 

—  escape  from  abdominal  operation  wound^ 

655 

from  inguinal  tumour,  656 

from  umbilicus,  656 

—  evacuation  in  enormous  numbers,  657 

—  expulsion  of,  drugs  for,  692 

—  female,  depositing  ova  in   liver,  689 

—  in  bile-ducts,  688,  689 

—  in   pulmonary  artery,   656 

—  in  upper   air-passages,  690 

danger  of,  691 

how  introduced,  690 

—  infection  by,  prophylaxis  against,  692 

—  invading  urinary  passages,  692 

—  invasion  of,  causing  liver  abscess,  690 

—  massive   accumulation   causing  occlusion 

of  intestine,  657 
Ascarince,  461 
Ascaris  causing  perforative  peritonitis,  65d 

—  characters,  461 

—  in   appendix   causing   intestinal   obstruc- 

tion, 654 

—  in  peritoneal   cavity,  656 

—  lacks   intermediate   host,   21 

—  lumbricoides ,     distribution     world-wide, 

463 

excretory  apparatus,  367 

expulsion   of,   754 

hosts  of,  464 

infection  by,  687 

experimental,  464,  465 

mode  of,  464,  465 

symptoms,  688 

injury  inflicted  by,  depends  on  num- 
ber in  host,  8,  9 

male,  hind  end,  371 

transverse  section  through  pos- 
terior extremity  of  body,  370 

method    of   keeping    alive,    754 

migration     from    small    intestine    ta 

other  parts  of  body,  464 


«4o 


'J  HE   ANIMAL   PARASITES    OF   MAN 


Ascaris    lumbricoides,    morphology,    463 

names  by  which  known  in  antiquity, 

464 

■ normal   habitat,    small    intestine,    464 

organs  of,  463 

—  —  ova    of,    463 

prevalence  in  young  children  in  tem- 
perate climates,  464 

self-infection  with,  on  part  of  experi- 
menter, 464,  465 

sites   of   body   invaded   by,   687,   688 

transverse     section     showing     organic, 

diagram  of,  362 

■ through,  diagram  showing,  364 

unrecognized    Dioctophyme    gigas    in 

man   traced  to,   431 

• —  maritima,   465 

- —  megalocephala,  nervous  system,  schema- 
tic representation,  365 

"tuft-like"  or  "  phagocytic  "  organs, 

362 

—  ova    of,    in    pus    in    case    of    abscess    of 

omentum,  657 

—  perforation   of  intestine  by,  656 
following    diseased    processes,    656 

—  sp.,  465 

—  texana,  465 

Ascites  and  abdominal  malignant  growth, 
Leydenia  gemmipara  associated  with, 
49,  50 

- —  association  of  Leydenia  ge7nmipara 
with,  49 

—  chylous,    from    Filaria    bancrofti    infec- 

tion, 678 

—  set  up  by  invasion  of  ova  of  Schistosoma 

japonicum,  282 
Ascitic    fluid    in    cultivation    of    Treponema 

pallidum,   125,   126 
Askanazy,  mode  of  infection  by  Opisthorchis 

felineiis,  254 
■ —  post-mortem   discoveries   cf    Opisthorchis 

felineus,  253 

—  Trichuris    trichiura,    420 

Asphyxia  following  invasion  of  upper  air- 
passages  by  Ascarides,  691 

Aspidogaster  attains  maturity  in  lower 
animals,   21 

Asses,   nagana   fatal   to,   94 

Atoxyl  and  antimony  combined  in  sleeping 
sickness,  622 

—  in  Indian  kala-azar,  626 

- —  in    infantile    kala-azar,    627 

—  in  sleeping  sickness,  622,  623 

—  Trypanosoma  rhodcsiense  resistant  to,  78 
Atractonema  gibbosum,  habitat,  4 

peculiar  characters  of  females,  5 

Auchmeromyia  (Bengalia)  depressa  as  cause 

of  myiasis  externa,  724 
"  larva  of  Natal,"  characters,  591 

—  luteola,  593,  594 

characters,  594 

geographical  distribution,  594 

larva  of  (Congo  floor  maggot),  593,  594 

how  destroyed,  594 

life-history,   614 


Auditory  meatus,  larvae  penetrating,  721 

external,    infected    with    Ehinospori- 

dium,   195,   196 
Austen,  ChironomidoR  described  by,  580 

—  description      of      larva      of      Ochromyia 

anthropophaga    (footnote),  590,   591 

—  myiasis   due  to  Sarcophaga,  590 
Autopsies  at  Tomsk,  human  parasites  most 

frequently   found   at,   253 

—  occTirrence  of  Linguatula  rhinaria  at,  526 


B. 


Babes,  A.,  cases  of  transmission  of  Demodex 
jolliculoruTii  canis  infection  to  man,  709 

—  Filaria  {?)  conjunctivoi  in  man,  405 
Babesia,    155,    172,    173,    174,    177 

—  bigemina,    geographical    distribution    of, 

177 

—  bovis,    causal    agent    of    "  Texas    fever " 

or  "  red-water  fever  "  in  cattle,  173 
cause  of  infectious  haemoglobiauria  in 

cattle,    177 
transmission  of,  by  Ixodes  riciniis,  177 

—  caballi,    177,    178 

cause  of  biliary  fever  in  equines,  177 

geographical  distribution  of,    177 

transmitting  agent,   178 

—  canis    and    B.    bovis,    life-cycle    in    tick, 

stages  of,   176,  177 

agents  of  transmission,   177 

cause  cf  malignant  jaundice  in  dogs, 

177 
cultivation  in  vitro  by  Bass's  method, 

172,   177 
life-cycle  in  infected  blood  of  dog,  175 

—  distribution   of  chromatin   in,    176 

—  ilivergens,   cause  of  European   red-water 

fever   in  cattle,   177 
geographical   distribution  of,   177 

—  morphology  and  hosts  of,  174 

—  muris,  morphology,  178 

—  nuclear  phenomena  in  species  of,  176 

—  ovis,  agent  of  transmission  of,  177 

cause  of  ''  carceag  "  in  sheep,  177 

geographical  distribution  of,   177 

transmitting    agent,    177 

—  parasites  of  red  blood  corpuscles  cf  mam- 

mals, 154 

—  pitheci,    178 

—  species    of,    177 

—  synonyms    (see    Piroplasma),    174 

—  tick  borne,  176 

development    in,    176 

Babesiasis,    symptoms    of,    178 

—  treatment    of,    178 

Baboon     serum,     action     on     Trypanosoma 

rhodesiense,  80 
Bacillus   lymphangiticus,   755 
Baelz,   prophylaxis  against  kedani,   703 
Baer,    C.    E.    von,    views    as    to    origin    of 

cercarise,  12 
Bagdad    sore,    parasite    of,    supposed    inter- 
mediate host,  575 
Bahr,   filariasis  in  Fiji,  401,  403 
Baker,  larvae  of  Aphiochceta  ferruginea,  583 


INDEX 


841 


Balantidiasis,  see  Dysentery,  balantidian  or 

ciliate 
Balantidium,   200 

—  coli,  200,  201 

dysentery    associated    Avith,    202,    203, 

637 

geographical   distribution,   201 

habitat  in  body,  201 

hosts    of,    7,    202 

in  man,  cases  recorded,  201 

morphology,    200 

transmission,   202 

—  giganteum,  see  Nyctotherus  giganteus 

—  minutum,  204 

habitat  in  body,  204 

in    diarrhoea,   204 

morphology,   204 

—  morphology,   200 

—  reproduction   of,   200 

Balbiani,  infection  by  Dioctojyhyme  gigas, 
432 

—  researches  on  silkworm  disease,  184 
Baleri,   causal   agent  of,   95 

Balfour,    A.,    coccoid   bodies   of    Treponema 

pallidiuu,    124,   125 
granules   in   digestive   tract  of  Argas 

persicus,  507 
Balfour,    A.,    and    Sambon,    researches    on 

Spirochceta  granulosa,  116 
Balsam  of  Peru,  application  in  scabies,  707 

•  —  in   crab   louse   infection,   712 

— in  head  louse  infection,  710 

■ ^  in  nasal  myiasis,  719 

—  permanent  m.ounting  agent  for  flukes,  471 
Balzer   and   Schimpff,   unusual   situation   of 

Sarcophaga  larvae,   723 
Barbagallo,    case    of    dermatitis    set    up    by 
Oxyuris   vermiciilaris ,  696 

—  method   of  evacuation   of   Oxyuridce,   697 
Barbeiro,   parasite   causing,   537 

Barbel   disease,  cause  of,   184 

Barkan,  nematode  in  human  eye,  412 

Barlow,   Craigia  hominis,  734 

Barth,  pseudomeningitis  following  infection 

by   Trichocephalus  dispar,  650 
Basile,  experiments  showing  that   infantile 

leishmaniasis  is  transmitted  by  lleas.  Ill 

—  transmission  of  canine  kala-azar  by  dog 

fleas,    103 

Bass,  C.  C,  cultivation  of  malarial  para- 
sites, 170 

Bass  and  Hall,  detection  of  ancylostome 
eggs,   473 

Bass  and  Johns  on  Entamoeba  buccalis,  43 

—  —  treatment   of   oral    endamoebiasis,   620 
Bass's  method,  cultivation  of  Babesia  (Piro- 

plasma)  canis  by,  in  vitro,   172,   177 

—  of  malarial  parasites,   170,  171,  172 

Bastianelli,  mosquitoes  in  relation  to  human 

malaria,   158 
Bat  parasites  (Streblidoe),  611 
Baths,  luke-warm,  in  trichinosis,  681 
3eattie    on    Rhinosporidium    from    Madras, 

197 
Becker,  trichocephalus  anaemia,  651 


Beclere,  method  of  extraction  of  Dracuncu- 

lus  medinensis,  390 
Bed  bug,  development  of  LeiA/j?» «nia  tropica 
in,   108 

probable     agent    of    transmission     of 

kala-azar,  107 

Texas  or  Mexican,  see  Conorhiniis  san- 

giiisuga 

see  also  Cimex  (Acanthia)  lectularia 

Bee   parasites   {Braulidoi),  611 

Bees,  microsporidiosis,  due  to  Nos.ema  apis, 

in,    185 
Beetle,  intermediate  host  of  Echinorhynchus 

moniliformis ,  478 
Begonin,  Oxyuris  in   appendix,  654 
Behrenroth,    prevention    and    treatment    of 

balantidian  dysentery,  G37 
Belascaris,  morphology,  463 
j    - —  cati,   morphology,   466 

! ovarian       tube,       transverse       section 

I  through,   369 

I transverse  section  through  head  part 

of,  466 
{    —  marginata,  morphology,  466 

Beneden,   van,  on   commensals,   6 
I    Bentley,     beta-naphthol     in     expulsion     of 
j  ancylostomes,   687 

Benzine  and  jsetroleum  in  crab  louse  infec- 
tion, 712 
- —  and  water  enemata  in  Trichuris  trichiura 
infections,  680 
t    —  enemata  in   arrest  of  trichinosis,  681 

—  high     injections     of,     in     evacuation     of 
I  Oxyuridce  to  be  avoided,  698 

I    —  inhalations  in  nasal  myiasis,  719 

Bergmann,  operation  results  of  cysticercus 
I  of  brain,  665 

—  Scolopendra  in  frontal  sinus,  721 
Berlin,    number    of    oxen,    sheep,    and    pigs 

slaughtered  in,   infected,   346 
of  pigs  found  trichinous  in,  430 

—  oxen  infected  with  Cysticercus  bovis  in, 

341 
Berti,  cause  of  ancylostome  anaemia,  648 
Bertramia,   194,   195 

Bertrand,    scolopendra    in    maxillarv    sinus, 
721 

i    Beta-naphthol  in  expulsion  of  ancylostomes, 

I  687 

j    —  ointment  in  copra  itch,  513 

Bete  rouge,  undescribed   species  of  Leptus, 

I  486 

j    Betten,     Caligus     curtus     invading     cornea 

I  (footnote),  483 

Big  game  reservoirs  of  nagana,  94 

of    Trypanosoma     rhodesiense,     69 

trypanosomes   innocuous  to,   70 

Bignami,  mosquitoes  in  relation   to  human 

malaria,   158 
Bile,  preservation  of  ova  of  flukes  in,  472 
Bile-ducts,    Ascarides    in,    688,    689 
in  results  of,  689,  690 

—  habitat  of  Clonorchis  sinensis,  259 

—  human,    thickened    and    dilated,    Amphi- 
merus  noverca  found  in,  258 

—  inhabited  by  Metorchis  truncatus,  262 


842 


THE   ANIMAL   PARASITES   OF   MAN 


Bile-diicts,   invasion   by  Clonorchis  sinensis, 
641 

—  pathological     changes     in,     set     up     by 

Clonorchis  endemicus,  260,  261 

—  and  liver,  habitat  of  Clonorchis  endemi- 

cus, 259,  260 
Bilharz,  discovery  of  Uymenolepis  nana,  323 

—  Porocephalus  constrictus ,  526 
Bilharzia  Mission,  report  of,  277 
Bilharziasis,   diagnosis,   643 

—  prognosis,    643 

—  prophylaxis  against,  644 

—  regions   of   body   affected    by,   642,   643 
— •  symptoms,   641  \ 

—  —  mainly   urinary,   641 

—  treatment,   643 

Biliary  fever  in  equines,  cause  of,  177 
Billings,   percentage   of   rats   infected   with 
trichinella,  427 

—  proportion  of  trichiuous  pigs  found  by, 

428 
Binotia,  characters,  565 
Bird  epithelioma  contagiosum,  207 
Birds,  blood  of,  Halteridium  parasites  occur 

in,  151 

—  developnjent  of  Plasmodium  relictum  in, 

170 

—  endoglobular   parasites,   similar   to  mala- 

rial, in,  discovery  of,  157 

—  experimental     infection     vi^ith     herpeto- 

monads,    739 

—  herpetomonads  in   blood  of,  739 

—  malaria  in,   spread  by  mosquito,   158 

—  mites    living   endoparasitically   in,   491 

—  sarcosporidia  in,   187 

—  species     of,    inhabited    by    Hymenolepis 

lanceolata,  329 
Bironella,   characters,   562,  570 
Bismuth  subnitrate  in  dysentery,  619 
Bitter  Koot  Valley  of   Montana,   mortality 

of  Rocky  Mountain  tick  fever  at,  504 
Blackhead  in  turkeys,  causal  agent,  145 
Blacklock,    experimental    host    of    Trypano- 
soma cruzi,  87 

—  and  Yorke,  Trypanosoma  equi,  98 

—  see  also   Yoi-ke  and  Blacklock 
Bladder,   amcebse   invading,   46 

—  means    of    access    of   Schistosoma    hcBma- 

tobium  to,  272 
- —  pathological  changes  in,  due  to  Schisto- 
soma hcematohium,  275 

—  worms,    development   of,    15 
explanation  of,  14 

Blaizot,   mode  of  transmission  of  relapsing 

fever,   120 
Blanchard,  Myriapoda  parasitic  in  intestine 

and  nose  of  man,  483 

—  on  Lamblia,   57,   60 

—  on   Monas   pyophila,   02 

—  on   nomenclature  of  amoebae,   31 
Blaps  mortisaga,  larva?  of,  in  stools,  542 
Blenorrhoea,    inclusion,    in    infants,    207 
Blepharitis  due  to  head  louse  infection,  710 
Blood,  changes  in,  in  ancylostomiasis,  683 

—  circulating,  morphology  of  Trypanosoma 

qamhiense  in,  73 


Blood,     citrated,     cultivation     method     for 
Leishmania  tropica,   108 

—  colourless,  of  insecta,   530 

—  corpuscle,  red,  number  of  malignant  ter- 

tian parasites  found  in  one,  167 

—  corpuscles,    red,    action    of    leucocytozoa 

on,    742 

attacked     by      quartan      malarial 

parasite,  not  altered  in  size  or  colour, 
166,  167 

— development     of     benign     tertian 

parasite    in,    160,    164,    165 

appearance    of    Schiiff- 

ner's  dots,   165,   166,   168 
malignant   tertian    parasite    in, 

appearance   of   Maurer's   dots,    168 

hsemogregarines  in,  153,   154 

— ingestion  by  amoebae,  35,  39,  42 

life-cycle  of  Nuttalia  equi  in,   173 

— separation   from   filaria  larvae,   395 

—  examination  of,  for  protozoa,  745 
in  diagnosis  of  trichinosis,  681 

—  films,  thick,  method  of  making,  747 

—  —  thin,   method   of   making,   747 

—  flagellates,  history  of,  67 
hosts   of,   67 

—  inoculation,  rinderpest  transmissible  by, 

742 

—  larvae  of  Loa   loa  in,  412,  414 

—  multiplication   of  trypanosomes   in,   71 

—  peripheral,  periodicity  of  larvae  of  Filaria 

bancrofti  in,  393,  394 

—  prevalence     of     filarial     disease     propor- 

tionate to  amount  of  Mikrofilaria  ban- 
crofti  in,   400 

—  protozoa  parasitic  in,  culture  media  for, 

744 

—  spirochaetes,  114 

—  supply   of   abdominal   and   pelvic  organs 

as   illustrating   distribution   of   Schisto- 
soma hcematobium  in  body,  272 

—  trypanosomes  in,  cultures  of,  69 
cyclical   variation,    78 

daily  number  from  case  of  Rhodesian 

sleeping  sickness,   79 

method  of  determining  number,  748 

periodicity  of,   69 

seasonal  variation,  69 

Blood-sucking  habit  of  Argas  persicus,  507 

of  Argas  reflexus,  506 

of  Cimex   lectularius   (bed  bug),  535 

of   Conorhinus   renggeri    (great   black 

bug   of   Pampas),   539 

— sanguisuga,  537 

of  Culicoides,  580 

of  fleas,  543 

of  Glossina  palpaUs,  607 

of  leeches,  701 

of  Leptidce,  603 

of   Linguatulidoi,   523 

of  mosquitoes  confined  to  females,  552 

of  Muscidce,  603 

—  —  of   Phlebotomus,   581 
! of  Pupipara,  611 

j    Blood-vessels,  migration  of  oncospheres  from 
I  intestine  to  liver  through,  302 


INDEX 


H: 


Blood-vessels,    Strong yloides    stercoralis    in, 

755 
Boas,  injection  of  emulsion  of  male  fern,  671 
Bodo,  63 
Bodonidoe,  61 

—  characters   of,   63 

—  genera   of,   63 

Body    lice,    human,    inhabited    by    Herpeto- 

monas  pediculi,  103 

prophylaxis  against,  615,  616 

see  also  Pediculus  vestimenti 

Bohland,   proteid   destruction   in    ancylosto- 

miasis,  647 
Boils  and  ulcers  due  to  invasion  by  larvae  of 

Cordylobia  anthropophaga,  592 

—  produced  by  Oestridce,  725 

Bo j anus,  views  as  to  origin  of  cercariae,  12 
Bollinger,  cases  of  Dipylidium  caninum  in- 
fection, 659 
Bolt,  sand  flies  in  North  China,  613 
Bond,  larvae  of  Muscidoe  in  nose,   720 
Bone-marrow,    development   of   crescents   of 
tertian  malignant  parasite  in,   169 

—  reu,  aamiuistration  in  Indian  kala-azar, 

626 
Bont  tick,  see  Amblyomma  hebrceum 
Boophilus  annulatus ,  transmission  of  Babe- 
sia bigejyiina  by,  177 

—  characters  of,  497 

—  species  of,  transmission  of  Babesia  bige- 

mina  by,  177 
Boracic  acid  fomentations  in  Oriental  sore, 

628 
Bordier,  Davainea  madagascarensis ,  662 
Borger,  cases  of  Ascarides  in  bile-ducts,  688 
Borrelia,   115 
Bosanquet,  molluscan  spirochaetes  breaking 

up  into  granules,  119 
Boschulte,  effects  of  bite  of  Argas  reflexus, 

506 
Bothriocephalus  anaemia,  644,  645 

cases  of,  645 

dissolution    of    parasitic    products    in 

serum  of  patients  with,  645 

experimental,  646 

Bouin-Duboscq  fluid,  749 

Bouin's  fluid,  749 

Brachycera   (flies),  characters,  582 

larvae  (maggots)  of,  parasitic  in  man, 

582 
Bradford,  Sir  J.  Rose,  see  Plimmer,  H.   O. 
Brain,  abscess  of,  set  up  by  amoebae,  35 

—  cysticerci  in,  335,  664,  665 

■ change  of  position,  665 

operation  for,  665 

percentage   of   cases,   664 

site,   664,  665 

symptoms,  665 

—  fourth  ventricle,  cysticerci  in,  symptoms, 

665 
treatment,   666 

—  paragonimiasis  of,  639 

diagnosis,  640 

prognosis,  640 

Brandesia  turgida,  host  of,  6 

Brandt,    effect   of   invasion    by   Dipylidium 
caninum  on  nervous  system,  649 


Braulidce    (bee  parasites),  611 

Braun,    developmental    cycle    of    Dibothrio- 

cephalus  latus,   16 
Brazil,  Trypanosoma  cruzi  prevalent  in,  83 
Breast,    tumour    of,    infection    by    Gnatho- 

stoma  siamense  associated  with,  385 
Breinl,  enlarged  glands  in  filariasis,  402 

—  researches  on  Spirochoeta  duttoni,  116 
Bremser,  origin  of  helminthes,  12 
Brieger,  effects  of  filmaron  oil,  672 
Britton,  fatal  case  of  myiasis  externa,  716 
Brock,  treatment  of  bilharziasis,  643 
Broden   and  Rodhain's  method   of  adminis- 
tering atoxyl  in  sleeping  sickness,  622 

Bronchi,  invasion  by  Paragonimus  ringeri, 

251 
Bronchitis  due  to  invasion  of  air  passages 

by  Paragonimus  ringeri,  251 

—  in  pigs  set  up  by  Metastrongylus  apri, 

433 

—  spirochaete   associated   with,    122,   632 
Bruce,    Sir    D.,    classification    of    trypano- 

somes,  72 

development  of  Trypanosoma  gam- 

biense  in  Glossina  palpalis,  74 

discovery  of  trypanosomes  in  blood 

of   horses   with   '*  nagana,"    68 

investigation  of  sleeping  sickness, 

68 

investigations      of       Trypanosoma 

rhodesiense  in  Glossina  morsitans,  82 

proportion  of  Glossina  palpalis  be- 
coming infected,  608 

question  of  distinction  or  identity 

of  Trypanosoma  brucei  and  T.  rhode- 
siense, 83 

tsetse-fly  transmitting  Trypano- 
soma rhodesiense,  608 

Zululand    strain    of    Trypanosom,a 

brucei,  94 

Brues  and  Sheppard,  insects  transmitting 
epidemic  poliomyelitis,  612 

Brumpt,  experimental  hosts  of  Trypano' 
soma  cruzi,  87 

—  on   Sergentella    hominis,    210 

—  subcutaneous  tumours  associated  with  in- 

vasion by  Onchocerca  volvulus,  418 

—  Tef^amitus  mesnili,  57,  624 

—  trichomonad  cysts,  56 

Brun's  symptom  of  cysticerci  in  fourth  ven- 
tricle, 666 

Biichholz,  tetanus  disappearing  after  expul- 
sion of  AscaridoB,  650 

Buff  coagulum  in  cultivation  of  Treponema 
pallidum,  126 

Bulgaria,  oestrid  larvae  in,  invading  human 
integument,  595 

Bunostomece,  456 

Burbot  (Lota  vulgaris),  muscles  of  trunk 
containing  plerocercoid,  313 

Burfield,  bilharziasis  of  appendix,  642 

Bursa  copulatrix  of  male  nematodes,  370 

Buschmucker,  mite   attacking  man,  486 

Biitschli,    O.,    on    Gregarines,    130 

on   Myxosporidia,   181 


53 


844 


THE   ANIMAL   PARASITES   OF   MAN 


C. 

CiECA,  intestinal,  of  nematodes,  364 

Caecum,  cysts  of,  oesophagostomum  con- 
tained in,  441,  443,  444 

Caffarena,  echinococcus  cysts  causing  urti- 
caria, 652 

Cairo,  fresh-water  molluscs  round,  cercariae 
of  bilharzia  type  in,  277 

Calandruccio,  experimental  infection  with 
Ascaris   lumbricoides ,  465 

self-infection    with    Oxyuris    vermicu- 

laris,  469 

—  on    experimental    amoebic    infection,    30 
Calcium   oxalate    crystals   in   endoplasm   of 

Lithocystis,  131 

—  salts,    internal    administration   for   cuta- 

neous and  muscular  cysticerci,  663 
Caligus   curtus  invading  cornea   (footnote), 

483 
Calkins  on  cultural  amoebae,  42 

—  on    genus    Craigia,    45 

—  on  vaccinia  and  variola,  208 
Calliobothrium,  larval  stage,  305 
Calomel  in  evacuation  of  Oxyuridce,  697 

—  in  flagellate  dysentery,  624,  625 

—  in  intestinal  myiasis,  728 

—  insufflations  of,  in  nasal  myiasis,  719 
following  cocainization  of  nose, 

720 
Calvertina,  characters,  562,  570 
Camels,  "  surra  "  in,  95 
Canaries,     susceptibility     to     infection     by 

Plasmodium   relictum,,   170 
Cancer,  association  with  invasion  by   Opis-    j 

thorchis  felineus,  254 

—  of  oral  cavity,  association  of  Entamoeba 

buccalis  with,  43 
Canestrini  and  Kramer,  species  of  Demodex, 

522 

of    Sarcoptes    enumerated    by,    518 

Canguary,  synonym-  of  Brazilian  trypanoso- 

iniasis,  87 
Canterbury   Cathedral,   Argas   reflexus   for- 
merly abundant  in,  506 
Cantlie,  J.,  sand  flies  biting  in  Hampshire, 

579 
Cape  ailment  (Port  Natal  sickness),  488 
Cappez,  nematodes  in  human  eye,  412 
Carbolic  acid  clearing  agent  for  flukes,  471 

injection   in   creeping   disease,   731 

Carbon  dioxide  snow,  application  in  Oriental 

sore,  628 
Carceag  in  sheep,  cause  of,  177 
Cardiac  form  of  Brazilian  trypanosomiasis, 

88 

—  stimulants  in  Asiatic  relapsing  fever,  631 
Caries,    dental,    association    of    Entam,oeba 

buccalis  with,  43 

kartulisi  with,  44 

Carini,  cysts  in  lungs  of  rats,  90 

—  phagedaenic    amoebae,    733 

—  treatment  of  espundia,  629 
Carlsbad  water  in  intestinal  myiasis,  728 
Carmine,  acetic-alum,  solution  of,  in  stain- 
ing flukes,  471 

—  solutions  of,  in  staining  flukes,  471 


Caryophyllaeus,  larvae  of,  305 

Casagrandi     and    Barbagallo,     Entamcebae 

31,    33 
Castellani,    A.,    demonstration    of   trypano- 

somes  in  cerebrospinal  fluid  from  sleep- 
ing sickness,  68 
experimental     production     of     copra 

itch,  513 

Nyctotherus    africanus,    206 

Toxoplasma  pyrogenes,  113 

Treponema   pertenue,    127 

and  Chalmers,  on  Ehinosporidium  in 

Ceylon,  197 
treatment  of  relapsing   fever,   630, 

631 

of  balantidian  dysentery,  637 

of  gangrenous  dysentery,  619 

of  Indian  kala-azar,  626 

of  infantile  kala-azar,  627 

of    malaria,    635 

of  Oriental  sore,  628 

— ■ —  of  sleeping  sickness,  622 

types  of  amoebic  dysentery,  618 

Castor  oil  in  evacuation  of  Oxyuridce,  697 

in  flagellate  dysentery,  625 

preliminary  administration  in  amoebic 

dysentery,  619 
Cat,  Dipylidium  caninum  parasitic  in,  322 

—  flea,  see  Ctenocephalus  felis 

—  host  of  Dibothriocephalus  latus,  313 
of  Paragonimus  kellecotti,  251 

—  infected       with       Dipylidium       caninum 

through  skin  parasites,  323 

—  intestine  of,  Isospora  bigemina  parasitic 

in,  149 

—  parasites    found    only    in,    6 

—  production  of  dysentery  in,  by  infection 

with  amoebae,  30,  35 

—  —  of  enteritis  in,  by  injection  of  amoebae 

into,  36,  37 

—  Tcenia  echinococcus  in,  356 

—  and  dog,  parasites  common  to,  6 
Catageiomyia,  characters,  563 
Cataphoresis  in  creeping  disease,  731 
Cataract,  ,  association      of     Filaria      oculi 

humani  with,  406 
Catarrh,  intestinal,  associated  with  Balan- 

tidium  coli,  201 
Caterpillars,  larvae  living  in,  discovery  of,  10 
Cathoemasia    (Distoma)    hians,    progeny   of, 

discovery,  12 
Cattle,    disease    in,    caused    by    Anaplasma 

organisms,   180 

—  East  Coast  fever  in,  cause  of,  178 
pathogenic  agent,  174 

—  European  red-water  fever  in,  cause  of,  177 

—  gall-sickness  in,  cause  of,  180 
supposed  causal  agent,  98 

—  infectious   haemoglobinuria  in,   cause  of, 

177 

—  migrations  of  larvae  of  Hypoderma  bovis 

in  body  of,  595 

—  nagana  prevalent  among,  and  generally 

fatal  to,  93,  94 

—  organs   infected   with   echinococcus,    per- 

centage of  frequency,  347 


INDEX 


84s 


Cattle,  red  dysentery  in,   cause  of,   147 

—  E-hodesian   fever  in,  carriers  of,  494 

—  "  surra  "   in,  95 

—  Texas  or  red-water  fever  in,  carrier  of, 

494 
causal  agent,  173 

—  Tryyanosoma  vivax  fatal  to,  99 
Caullery    and    Mesnil,    Actinomyxidia,    187 
Haplosporidia,    194 

Caval  system,  means  of  access  of  Schisto- 
soma hcematohium  to,  272 

Schistosoma  hoematobium  in,   274 

Cedar-wood  oil,  mounting  agent  for  flukes, 
471 

Cell    inclusions,    207,    208 

Celli,  discovery  of  movements  in  malarial 
parasites,   157 

—  and    Fiocca,    species    of    amoeba    distin- 

guished by,   31 
Cellia,  characters,  562,  569 
Centrorhyncus,  581 

Cephalina,  morphology  and  hosts  of,   135 
Cephalodiscus  nigrescens,  195 
Ceratixodes,  characters  of,  497 

—  hosts  of,  497 
Ceratomyxa,  184 
Ceratophyllus  anisus,  548 

—  distinctive   characters,   545 

—  fasciatus,  carrier  of  plague  bacillus,  543, 

547 
development    of    crithidial    forms    of 

Trypanosoma  lewisi  in  rectum  of,  91,  93 
transmission    of    Trypanosoma    lewisi 

by,  88 

—  londiniensis ,   548 

Ceratopogonince  (midges),  characters  of,  580 

—  geographical  distribution,  580,  581 
Cercaria  bilharzia,  characters,  754 

— ■  bilharziella,  characters,  754 
Cercarise,     bilharzia    type,     in     fresh-water 
molluscs  round  Cairo,  277 

—  (larval  stages)  of  Trematodes,  225,  227, 

228 

—  origin  of,  early  views  as  to,   12 
Cercomonadidce,  61 
Cercomonads,  encysted  stages  of,  62 

—  presence  in  intestine,  62 
Cercomonas,    characters,    61,    736 

—  hominis,   61,    736 

characters,  61 

flagellum   of,   61,   62 

—  intestinalis ,  characters  of,  54 

—  longicauda,  736 

—  parva,  737 

—  vaginalis,  62 

Cereals,  mites  infesting,  effects  on  man,  489 
Cerebrospinal   fluid  from   cases  of   sleeping 

sickness,  trypanosomes  in,  68 
Cesspools,  screening  against  mosquitoes,  636 
Cestoda,  animal  nature  of,  always  known, 

282 

—  cirrus  pouch,  295 

—  cortical  layer,  289 

—  development  of,  13 

—  developmental  cycle  of,  discovery,  16 
direct,  17 


Cestoda,   egg-shell    substance,   297 

—  eggs  of,   297 

—  excretory  apparatus,  291 

vessels,    collecting   tubes,    291 

frontal  anastomosis,  291 

island   formation,   292 

—  fixation,  staining  and  clearing  of,  472 

—  genital    apparatus,   293 

female,  295 

male,  293,  294 

papilla,   295 

—  infection  by,  644 

—  life  spent  in  intermediate  and  final  host, 

18 

—  loss  of  digestive   system  in,  3 

—  medullary  layer,  289 

—  myoblasts,   289 

—  nervous  system  of,  289,  290 

—  of  man,  309 

classification,  308 

relative    frequency,    342 

—  ovaries,  295 

—  oviduct,  295 

—  —  course  and  situation,  295,  296 

—  parauterine   organs   or  uterine   capsules, 

297 

—  preservation  and  examination  of,  472 

—  relation  to  Turbellaria,  19 

—  rostellum,   289 

—  segmented  and  single-jointed  connection, 

20 

—  sexual  orifice,  male,  295 

—  suckers,  289 

—  testes,  293 

—  uterus  of,  296 

parenchymal   capsules,   296 

—  vagina,  295 

—  vas  deferens,  294 

Cestode  tuberculosis,  acute,  303 
Cestodes,  see  Cestoda  (above) 
Chagas,     morphology     and     life-history     of 
Trypanosoma  cruzi,  84-87 

—  multiplication   of   Trypanosoma   cruzi  in 

vertebrate   host,   85 

—  on    Cercomonas    parva,    737 

—  researches  on  Brazilian  trypanosomiasis, 

87,  88 

—  reservoir  of  Trypanosoma  cruzi,  87 

—  schizotrypanosomiasis,  88 

—  Trypanosoma  cruzi,  83 
Chagasia,  characters,  562,   570 

Chagas's  disease,  see  Trypanosomiasis , 
Brazilian 

Chalazion  due  to  infection  by  Demodex  fol- 
liculorum,  708 

Chalmers  and  O'Farrell,  treatment  of 
bronchial  spirochaetosis,  633 

see  also  Castellani  and  Chalmers 

Chatin,  Metastrongylus  apri  in  human  in- 
testine, 433 

Chatterjee,  Pentatrichomonas  bengalensis, 
624,    735 

Chatton  and  Lalung-Bonnaire,  entamoebae  of 
vertebrates,  34 

Cheese-fly,  583 


84^ 


THE   ANIMAL    PARASITES   OF   MAN 


Chelifer  cancroides,   pseudoparasite   in  man 

(footnote),  484 
Cheyletus,  characters,  516 

—  mericourti,  517 

CJiiari,  case  of  Oxyuridce  in  nose,  696 
Ciiiastopsylla,   distinctive   characters,   545 
Chicken  cholera,  possible  carrier  of,  579 
Chigoe,  see  DermatophiUis  penetrans 
Children,    administration    of    male    fern   to, 
671,  672 

—  Dipylidium  caninum  infection  prevalent 

in,  322,  659,  660 

—  enteritis  verminosa  in,  688 

—  flagellate  dysentery  in,  56 

— ■  invasion  by  Hymenolepis  nana,  324,  661 

—  lymphangitic    attacks    in,    from    Filaria 

bancrofti  infection,  676 

—  native,  carriers  of  malarial  parasites,  636 

—  —  latent  malaria   in,   158 

—  oxyuriasis  in,  695 
treatment,  697,  698 

—  young,     prevalence     of    Ascaris     lumbri- 

coides  in,  in  temperate  climates,  464 

Chilodon  dentatus,  206 

- —  uncinatus,  206 

Chilodon-like  organisms  found  in  gonor- 
rhoea,  206 

Chilomastix  and  Tetramitus,  differential 
characters,  735,  736 

—  (Tetramitus)  mesnili,  57,  735 
—  diagrams   of,   736 

China,   North,   sand   flies   and   fever  due   to 

them   in,   613 
Chironomidm         (midges),        distinguishing 

characters   from    mosquitoes,   579 

—  —  larvae   of,   579 
Chironomus,  wing  of,  579 
Chlamydophrys,  47,  759 

—  characters  of,   47 

—  enchelys,  asexual  multiplication  in  faeces, 

47 

characters  of,  47 

^  —  encysted  form,  48,  49 

free  motile  form  and  dividing  organ- 
isms, 47,  48 

habitat,  47 

plasmogamic  union,  48 

—  —  sexual  multiplication,  48 

—  stercorea,  47 

see   also   Chlamydophrys  enchelys 

Chlamydozoa,   207,   210 

—  characteristic  features,  208 

—  granules  of,  filtration  experiments  with, 

209 

—  mode  of  division,  209 

—  possible    cultivation,    210 

—  relationship  of,  210 

—  systematic    position,    210 

—  tissues  inhabited  by,  209 
Chlamydozoon,   life-history   of,   209 
Chloroform,  administration  in  expulsion  of 

ancylostomes,  686,  687 

—  as  tapeworm  drug,   674 

—  hypodermic  injection  in  creeping  disease, 

731 

—  in  t>xpulsion  of  guinea-worm,  676 


Chloroform     injections     or     inhalations     in 

nasal   myiasis,   720 
Chloromyxiace,  184 

Chloromyxum  leydigi,  trophozoite  of,  182 
Choanoflagellata,  52 
Cholera  Investigation  Commission,  work  of, 

with  regard  to  dysentery,  30 

—  motions,  spirochaetes  found  in,  122 

—  spread   by   house-fly,   586 
Cholodkowsky,   "  wormlet  "   burrowing   into 

human   epidermis,   599 
Chorea,    case    of,    cured    after    expulsion    of 

Taenia,  648 
Chorion  enveloping  ova  of  nematodes,  371 
Chorioptes  bovis,  521 

—  characters,   517 

—  species  found  on  man,  521 
Christophers,    on    Babesia    cants,    176,    177 

—  on    Lencocytogregarina    canis,    155 

—  see   Stephens   and   Christophers 
Christopherson,    J.    B.,    case    of    espundia, 

108,  628 
Chrithoptes  nionunguiculosiis,  489 
Chromidial  apparatus  of  protozoa,  26 
Chrysoconops,  577 

Chrysomela  hoemoptera,  gregarine  from,  131 
Chrysomyia  and  Pycnosoma,  distinguishing 

features,  588 

—  (Cotnpsomyia)     macellaria     (screw-worm 

fly),  587 
larvae  of,   invasion   by,   fatal 

results  from,  587 
—  regions    of    body    invaded 

by,  587 

references  to,   587 

synonyms,    587 

—  viridula,  characters  of,  588 

larvae   (maggots)  of,  discharged  from 

nose,  588 
Chrysops  host  of  Filaria  loa,  601 

—  dimidiata,  601 

—  silacea,   601 

—  transmission  of  surra  by,  601 
Chrystia,    characters,    562,   568 
Chyluria,  association  of  Hymenolepis  mada- 

gascarensis  with,  662 

—  following      infection      by      Eustrongylus 

gigas,   683 
by  Hymenolepis  nana,  661 

—  from  Filaria  bancrofti  infection,  677 
treatment,  677 

—  in  filariasis,  402 

—  without   lymphatic    obstruction,    401 
Cigarettes,     paper,    smoking    of,    in    nasal 

myiasis,  719 
Ciliata,    198 

—  classification,    199 

—  macro-nucleus  and  micro-nucleus  of,  198 

—  morphology  of,  198 

—  peristome  of,   198 

—  reproduction  of,    198 
Ciliophora,    198 

Cimex   (Acanthia)  lectularia  (bed  bug),  see 
Cimex    lectularius    (following) 

—  honeti,  host  of  Trypanosoma  criizi,  87 

—  ciliatus,   537 


INDEX 


847 


Cimex  columbarius ,  536 
bite  of,  536 

—  hirundinis  (swallow  bug),  537 

—  lectularius    (bed  hug),   artificial   host   of 

Trypanosoma  criizi,  87 

bite  of,  treatment,  713 

■  —  blood-sucking,    535 

— —  characters   and   habits  534 

extermination  of,  713 

infection  by,  713 

diagnosis,  713 

larval  stage,  535 

ova  of,  535 

peculiar  odour  emitted  by,  535 

—  — persistent      accompaniment      of 

man   by,   535 
possible    transmission    of    kala- 

azar   by,    107,    713 
transmission    of   SpirochcBta   re- 

currentis    by,    120,    121 

—  macrocephalus ,  536 

—  rotundatus    (tropical   bed  bug),   535,   536 
carrier    of     virus     of    polio- 
myelitis, 536 

geographical        distribution, 

536 
points  of  distinction  from  C. 

lectularius ,  536 
—  possible       connection       with 

kala-azar,   536 

—  sp.,  connection  with   Oriental  sore,  536 
CimicidcB,  characters,   534 

Circulatory     system,     disturbances     of,     in 

ancylostomiasis,  683 
Cisterns,  screening  against  mosquitoes,  636 
Cladorchiina>,  231,  234 

—  male   and   female  organs,   234 

—  morphology,  234 

Clark,  see  Howard  and  Clark 

Clasping  and  clinging  organs  in  permanent 
parasites,  4 

Climates,  temperate,  prevalence  of  Ascaris 
lumbricoides  in  young  children  in,  464 

Clonorchis  endemicus,  geographical  distri- 
bution, 260 

habitat  and  hosts  of,  259 

intermediate   host  of,  first  unknown, 

261 

second,  261 

life-history,   261 

—  —  morphology,  259 
synonyms,   259 

—  morphology,  258 

—  sinensis,  geographical  distribution,  259 

—  —  habitat  and  hosts  of,  259 

infection  by,  640 

diagnosis,  641 

prophylaxis  against,  641 

—  —  —  syniTotoms,  640,  641 

treatment,  symptomatic,  641 

morphology,  258,  259 

organs  of,  diagram  showing,  259 

ova   of,   259 

sites  of  invasion  in  body,  640 

Cloquet,  destruction  of  eyes  by  Sarcophaga 
larvae,  723 


Clothes  louse,  see  Pediculus  vestimenti 
Cnidosporidia,    129,    194 
Cobbold,  Ligula  niansoni,  318 
Cocaine,    application,    followed    by    calomel 
insufflations,  in  nasal  myiasis,  720 

—  hypodermic  injection  in  creeping  disease, 

731 
Coccidce  (scale  insects),  532 
Coccidia,  copulation  in,  137 

—  experimental  infection  with,  136 

—  ova    of    helminthes    mistaken    for    (foot- 

note),  137 

—  pathogenicity   of,   136 

Coccidia-like  organisms  in   various  diseases 

of  man,  150 
Coccidiidea,    129,   135,   137 

—  characters  and  habitat,  28 

—  classification   of,    141 

—  gametes  of,  137,  139,  140. 

—  hosts  of,   137 

—  life-cycle  of,  138-141 

—  macrogametes,    139,    140 

—  merozoites  of,   138,   139,   140 

—  microgametes  of,  137,  139,  140 

—  morphology  of,    138 

—  occurrence,    137 

—  oocysts  of,   141 

—  schizogony   in,   138 

—  sporoblasts  and  sporocyst  of,  141 

—  sporozoites  of,   138,  139,  140 
Coccidioides  imniitis,   150 

—  pyogenes,   150 
Coccidiomorpha,    129,   151 
Coccidiosis.    avian,    142-145 

—  diagnosis    of,    742 

—  human,  doubtful  cases,  149 

hepatic  cases,  148 

intestinal    cases,    148 

—  in    cattle,    147,   741 

—  in   rabbits,    intestinal    and   hepatic,    145, 

147 

—  rinderpest  mistaken  for,  741 
Coccidium  jalinum,   150 
hosts  of,   150 

Cochin-China       diarrhoea,       see       Diarrhoea 

(Cochin-China) 
Cockchafer,    intermediate    host    of    Echino- 

rhynchus  qigas,  477 
Cocoons  of   Hirudinea,   481 
Cod-liver   oil,    inunctions   of,   in   evacuation 

of  Oxyuridm,  698 
Coelosporidiuni,  195 
Coenurus,  definition  of,  301 

—  cerebralis,  experimental  rearing  of  tape- 

worms from,  15 

reared  experimentally,  15 

.  section     showing     cephalic     invagina- 
tions, 304 

—  scolices  in,  303 
Coleoptera,  characters,  531,  532 

—  larvae  of,   accidental   parasites,   542 
Colitis  associated  with  B alantidiiim  coli,  201 
with   intestinal    amoebae,  30 

—  mucous,  complicating  intestinal  myiasis, 

726,  727 
Collargol  in  balantidian  dysentery,  637 


848 


THE   ANIMAL   PARASITES    OF   MAN 


Collargol,    irrigation    of    lower    bowel    with, 
in   gangrenous   dysentery,   619 

—  rectal  administration  in  billiarziasis,  643 
Colon,  cysts  of,  (Esophagostoniiim  contained 

in,  441,  443,  444 

—  descending,   means  of   access   of  Schisto- 

stoma  hceynatobium  to,  272 
Colorada,  mite  attacking  man,  486 
Colpoda  cucullus,  204 
Comedones,  removal  in  infection  by  Demo- 

dex  folliculorum ,  708 
Commensals,  6,  20 

—  nature  of,  6 

Congo  floor  maggot,  593,  594 
Conjunctiva,  dipterous  larvae  in,   716 

—  trachoma    bodies    in    infected    epithelial 

cells  of,  209 
Conjunctivitis  due  to  head  louse  infection, 

710 
Connective  tissue,  subcutaneous  invasion  by 

Loa  loa,  678 
Conorhinus    megistus    intermediate   host   of 

Trypanosoma  cruzi,   83,   537 
see  Triatoma  megista 

—  nigrovarius ,  bite  of,  539 

—  protractus ,  539 

—  renggeri    (great   black   bug  of   Pampas), 

blood-sucking,  539 
- —  rubrofasciatus,  bite  of,  538 
trypanosome  inhabiting,  99 

—  sanguisuga     (blood-sucking    cone    nose), 

bite  of,  537,  538 
ova  of,  538 

—  sp.  (?),  bite  of,  symptoms  following,  538 

—  variegatus,  539 

Conseil,  mode  of  transmission  of  relapsing 

fever,  120 
Constipation,    prevention    during    malarial 

attacks,  635 

—  set  up  by  Ascarides,  657 
Copaiva  balsam  in  bilharziasis,  643 
Copper,  black  oxide  of,  as  tapeworm  driig, 

674 
Copra  itch,  mite  causing,  513 

—  —  treatment.    513 

Copulation,   modes  of,   in   tapeworms,   297 
Cordylobia  anthropophaga,  characters,  592 

geographical  distribution,  593 

hosts  of,   592 

larvae  of,  characters,  592 

lesions  set  up  by  invasion,  592 

life-history,   592 

references  to,  593 

—  grunbergi,  larvae,  characters,  591,  592 
synonyms,   591 

—  rodhani,  593 

geographical  distribution,   593 

Coreotrypanosis,  87 

Corethra,  565 

Corethrina,  565 

Coriscus   subcoleoptratus,   bite   of,   540 

geographical  distribution,  540 

synonyms,  540 

Cornea,   parasitic   crustacean    (Caligiis    cur- 

tus)  invading,  483 
Cortical  layer  of  Cestoda,  289 


Councilman    and    Lafieur    on    nomenclature 

of  amoebae,  31 
Couret  and  Walker,  J.,  culture  medium  for 

intestinal  amoebae,  743,  744 
Couvy,  Spirochceta  gallinarum,  119 

—  see  Marchoux  and  Couvy 
Cover-slip  preparations,  748 

Cows  attacked  by  Leptus  autuninalis,  486 

Crab  louse,  see  Phthirius  inguinalis 

Craig,  C.  F.,  on  Entarnceba  histolytica,  37, 

41 
on    Paramoeba    (Craigla)    hominis,   44, 

45 
Craigia   hominis,   45,   734 

—  migrans,  734 
Craigiasis,  734 

—  nature   of,   734 

Crane-fly,    Gregarina    longa    from    larva   of, 

130       • 
Craw-craw    (filaria    infection    of    the    skin), 

378,    514 
Crawley,  experimental  infection  with  Sarco- 

cystis,    192 

—  movements  of  gregarines,  131 
Creeping  disease,  cases  of,  various  authors 

reporting,   729 

—  —  clinical  symptoms,   730 

—  —  duration,  731 

localization  of,  730 

mode  of  origin,  729,  730 

synonyms,    729 

■  treatment,  731 

methods      reported       by       various 

authors,   731 

—  eruptions,   599 

Creosote  in  bronchial  spirochaetosis,  633 

—  mounting   agent   for   flukes,   471 
Creplin,  discovery  of  progeny  of  Diphyllo- 

botlirium    {Bothriocephaliis)    ditremnm, 
13 

—  psorosperms,    181 

Cristina   and   Caronia,   treatment   of   infan- 
tile kala-azar,  627 
Crithidia,  67,   103 

—  inoculation    experiments   with,    104,    738 

—  fasciculata,  104 
host  of,  104 

—  qerridis,  738 

—  hosts  of,  104 

—  hyalommm,  104 

—  melophagia,  host  of,   104 

—  morphology,  104 

—  natural  flagellates  of  Arthropoda,  104 
Crustacea,  parasitic,  change  of  original  fea- 
tures in,  4 

loss  of  digestive  sy^stem  in,  3 

or  free-living,  invading  man  abnor- 
mally  (footnote),  483 

Csokor,  mode  of  infection  in  intestinal 
myiasis,  727 

Ctenocephalus  canis,  herpetomonad  inhabit- 
ing, 103 

transmission    of    Trypanosoma    lewisi 

by,  88,  90,  92 

—  distinctive  characters,  545 

—  felis  (cat  flea),  547 


INDEX 


849 


Ctenocephalus,  hosts  of,  547 
Ctenophthalmus,       distinctive       characters, 

545 
Ctenopsylla,  548 

—  distinctive  characters,  545 

—  musculi,    transmission    of    Trypanosoma 

lewisi  by,  90 

Cucumber  seeds  in   intestinal  myiasis,  728 

Culex  and  Anopheles,  larvae  of,  position  in 
water  compared,  554 

— •  —  ova  of,  method  of  depositing  com- 
pared,  554 

points  of  difference  between,   551 

—  characters,   564 

—  fatigans     (common    tropical    gnat),    dis- 

tinguishing character,  576 
transmission  of  Filarise  by,  576 

—  head  of  male  and  female,  549,  556 

—  human    malaria    not    spread    by    species 

of,    158 

—  larva  of,  553 

—  ova  of,  557,  558 

—  pipiens    (common    gnat),   575 

—  —  characters,  576 

ova  of,  localities  selected  for  deposi- 
tion, 553 

—  species   of,    development    of    Plasmodium 

relicUim  in,  170 
Ctilicidoe,   classification   of,   501 

—  number  of  species,  552  , 

—  scales   of,   560 

CulicincB,   characters,    563,   571 
Culicoides,   blooi-sucking  habits   of,   580 

—  larvae  of,  580 

—  ornatus,  bite  of,  581 

—  possible   carrier   of   germ    of   Delhi   boil, 

580 

—  pupae  of,  580 

Culture   media   for   amoebae,   742 

—  —  for    blood    protozoa,    744 
Cunningham,        discovery       of       intestinal 

amoebae,  29 
Cutaneous     glands,     unicellular,     of    nema- 
todes, 361 

—  tumours  due  to  cysticerci,  characteristics 

of,  662,  663 

Cyclasterium,    208 

Cyelocoelum  mutabile,  progeny  of,  dis- 
covery,   12 

Cycloleppteron,  characters,  561,  567 

Cyclophyllidea,   308 

Cyclopidce,  characters,  390 

Cyclops,  characters,  390 

—  intermediate  host  of  Draciinculus  medin- 

ensis,  388 

—  virescens,  389 
Cyclospora,  141 

Cyrtoneura  stabulans,  larvae  of,  habitat,  585 
Cysticerci,    cutaneous    and    muscular,    (sym- 
ptoms   set  up  by,  663 
treatment,  663 

—  development  from  oncosphere  of  Teen  iidcB, 

303 

—  early  researches  on,  282 

—  experimentally    reared    from   tapeworms, 

15 


Cysticerci,  number  of,  in  relation  to  species 
of  Taeniae,  16 

—  origin  of,   14 

—  regions  of  body  site  of,  663,  664 

—  subretinal,  664 

—  tapeworms    experimentally    reared    from, 

15 

Cysticercoid,  morphology  of,  diagram  show- 
ing, 301 

Cysticercoids,  echinococcus-like  conditions 
in,  304 

Cysticercus  acanthotrias ,  336 

and  C.  cellulosoi,  336,  337 

—  hovis,   340 

amount     of     prevalence     in     ox,     340, 

in   Prussia   and  Berlin,   341 

, artificial    infection    of   human    beings 

with,  340 
rarity  in  man,  341 

—  celluloscR,  amount  of  injury  inflicted  by, 

depends  on  situation  in  body,  8 

development  to  ToBuia  solium,  340 

infection    of    skin    and    subcutaneous 

tissues,  662 

—  —  and   C.   acanthotrias,  336,   337 
decrease    of    frequency    in    pork,    how 

effected,  334 

development,   time   taken   for,   334 

habitat,  332,  333 

—  -  liosts  of,  332 

—  —    how  conveyed  to  man,  334,  335 
'u  man,  335 

I    -—  long  persistence  of,  337 

i in  sheep,  337 

organs  of  body  invaded  by,  335 

sex  distribution  of  invasion  by,  335 

vitality  of,  334 

—  development   of,   301,   302 
diagram  showing,  303 

:    —  fasciolaris,  host  of,  338 

' possible  means  of  spread  to  man,  338 

j    —  forms  of,  301 

!    —  morphology  of,  diagram  showing,  301 
'    —  avis,  337 

1    —  papilliform    invagination    into    bladder, 
301 

—  pisiformis,  hosts  of,  338 

in  evaginated  condition,  304 

— ■  racemosus ,  335,  336 

—  tenuicoUis,  337,  338 
experimental    rearing    of    tapeworms 

from,    15 

—  with  developed   scolex  at  bottom  of  in- 
vagination,   304 

Cystoflagellata,  52 

Cysts,     intestinal,     CEsophagostomum     con- 
tained in,   441,  443,  444 
Cytorhyctes,  208 

—  aphtharum,  208 

—  cytoplasm  of,  209 

—  luis,   124,  208 

—  scarlatince,  208 

—  vaccinice,  208 

—  variolce,  208 
minute  granules  in,  210 


850 


THE    ANIMAL    PARASITES    OF    MAN 


Dactylomyia,  characters,  562 
Daniels,   C.    W.,   iridocyclitis   in   trypanoso- 
miasis,  623 

—  —  parasitic     coleopterous     larva      (foot- 

note),   542 

treatment   of  trypanosomiasis,   622 

yellow   pigment   in   kidney    and   liver 

cells   in    ancylostomiasis,   647 

Danielsia,  characters,  564 

Danilewsky,  discovery  of  endoglobular  para- 
sites, similar  to  malarial,  in  birds,  157 

of  Leucocytozoa  by,  153 

Danube,  banks  of,  Simulium  columbaschen- 
sis  plague  on,  578 

Darling,  experimental  infection  with  Sarco- 
cystis   muris,    192 

—  Endotrypanum    schaudinni,    99 

—  Histoplasma  capsulatum,   112 

—  researches    on    Entamceha   tetragena,   38, 

40,    41 
Darwin,  Charles,  great  black  bug  of  Pam- 
pas (Conorhinus  renggeri),  539 
Dauernheim,  Ascarides  in  bile-ducts,  688 
Daughter  cysts  of  echiuococcus,  350,   351 

mode   of   origin,   350,   351,   352 

Davaine,   Cercomonas   hominis,   61 

—  milk  cure  in  expulsion  of  Strong yloides 

stercoralis,  675 

—  mode  of  development  of  Ascaris  lumbri- 

coides,  464,  465 
Davainea  asiatica.  662 
morphology,  330 

—  madagascarensis ,   association   with   chyl- 

uria,  662 

cases   of  human   infection   by,   330 

morphology,   329 

—  morphology,  329 
Davaineidce,  309,  329 
Davaineinoi,  309,  329 

Davidson,  bite  of  Rasahus  biguttatus ,  540 

Deeks,  W.  E.,  treatment  of  amoebic  dysen- 
tery,  619 

Deguy,  see  Labadie-Lagrave  and  Deguy 

Deinocerites,  characters,  564 

Delanoe,  pneumocysts  in  rats,  90 

Delhi  boil,  see  Oriental  sore 

Demarquay,  observation  of  Filaria  ban- 
crofti  in  man,   390 

Demodex  folliculorum,  521 

affecting  eyelids,  708 

as   cause   of   chalazion,   708 

characters,  522 

infection  by,  treatment,  708 

views  of  different  authors  respect- 
ing,  708 

synonyms,   522 

var.    canis,   522 

infection     by,     transmission     from 

dog   to   man,   709 

treatment,   709 

Demodicidoe  (mites  of  hair  follicles),  charac- 
ters, 522 

Dendriomyia,  characters,  565 


Dengue    fever,    micrococcus    supposed    car- 
rier,  576 
Dermacentor,  characters  of,  497 

—  occidentalis ,  bite  of,  effects,  504 

characters  and  morphology,  504 

(wood  tick),  characters,  504 

—  reticnlatus,   hosts  of,  502,  503 

transmission  of  Babesia  caballi  by,  178 

—  variabilis,   505 

—  venustus,  characters  and  morphology,  503 

hosts  of,  wild  and  domestic,  504 

Dermanyssus   gallinoe,  disinfection  methods 

against,  704 

effect  on  skin  of  host,  492 

infection,  symptoms  set  up  by,  703 

morphology,  492 

synonyms,  492 

—  hirundinis,  disinfection  against,  704 

skin   affections  due  to,  492 

Dermatitis  intertriginoides  set  up  by  Oxyii- 

ris  vermicularis ,  696 
Dermatobia  cyaniventris,  596,  597 

characters,   598 

geographical   distribution,  598 

hosts  of,  596 

larvae  of,  characters,  597 

local   or  vernacular  names   for,   598 

see   also  Mosquito  worm 

—  noxialis,  597 

larvae  of,  skin  disease  caused  by,  725 

Dermatocentor     reticulatus,     var.     occiden- 

I  talis,  carrier  of  Rocky  Mountain  spotted 

I  fever,  496 

I    Dermatodectes,   521 

!    Dermatophagoides   scheretnetewskyi,   521 
Dermatophilus,   543 

—  ccBcata,  544 

—  penetrans     (jigger,     chigoe),     characters 

and  morphology,  544 

geographical    distribution,    544 

j    —  (Sarcopsylla)   penetrans    (sand   flea),   en- 
I  trance  beneath  skin,  714 

1 host  of,  613 

'    — lesions  produced  by,  714 

I    — treatment,  715 

I    — possible  carrier  of  leprosy,  613 

Dermo-muscular  layer  of  nematodes,  361 
Derrieu    and    Raynaud,    chronic    dysentery 
due    to    trichomonads,    624 

trichomonad-like  organism  discovered 

by,   624,   735 
Desvoidea,  characters,  563 
Deve,   experimental   development   of  hyatid 

scolices,   353 
Diarrhoea  associated  with  Balantidium  coli, 
201 

with  invasion  by  Balantidium  minu- 

tum,  204 

with    Lamblia   intestinalis ,    59,    625 

with    Trichomonas    hominis,    54,    624 

with   presence   of    Watsonius   watsoni, 

235 
with  Prowazekia  asiatica,  65 

—  blood-stained,    in    Strongyloides    sterco- 

ralis infection,  674,  675 


INDEX 


851 


Diarrhoea   caused    by    Difdmus    tunensis,    57 

—  chylous,    from    Filaria    bancrofti    infec- 

tion, 678 

—  (Cochin-China)  in  cases  of  infection  with 

Strong yloides  stercoralis,  380,  381 

—  flagellate,    623 

— ■  climatic  distribution,  623 

—  infantile,  spread  by  house-fly,  586 

—  white   (or  white  scour),  in  fowls,  causal 

agent,   145 
DibothriocephalidcB,  308,   309 

—  morphology,  308 
Dibothriocephalince,    308,    309 

—  morphology,  308 
Dibothriocephalus,   309 

—  cordatus,  315 

cephalic  end,  315 

hosts  of,  315 

—  excretory     apparatus,     collecting     tubes, 

island  formation,  292 

—  felis,  313 

—  latus,  310 

chains  of  segments,  311 

development,   311 

developmental  cycle  of,  16 

disturbances  produced  by,  in  man,  314 

duration  of  life,  315 

embryophore  of,  298 

experimental   infection   of   man   with, 

312 

geographical   distribution,   313,  314 

growth  of,   306 

—  —  habitat  in  man,  658 

head  of  transverse  section,  311 

hosts  of,  313 

human  parasite  invading  animals,  7 

in  muscles  of  trunk  of  burbot,  313 

intermediate   host  of,   255 

means    of    transmission    to    man    and 

other   hosts,   314 

mode   of  infection,   658 

morphology,  310 

ova  of  Fasciola  hepatica  to  be  dis- 
tinguished from,  242 

ovum  of,  development,  312 

parasitic      association      with      Tcsnia 

solium,   658 

percentage  in  sufferers  from  tape- 
worms in  various  localities,  314 

plerocercoids  of,  313 

habitat  and  host,  311 

how  destroyed,  315 

inhabiting  fish,  314 

proglottids  of,  average  number  found 

daily,  312 

proglottis,  fairly  mature,  stained  pre- 
paration,  311 

prophylaxis  against,  668 

supposed   origin   of,   11 

— -  —  symptoms  produced  by  infection  by, 
667,   668 

synonyms,  310 

topographical      anatomy,      transverse 

section  through  proglottis  showing,  296 

—  morphology,  309 

—  parvus,  habitat,  316 


Dibothriocephalus  parvus,  how  distin- 
guished from  D.  latus  and  D.  cordatus, 
316 

morphology,  316 

—  plerocercoid  of,   300 

—  synonyms,   309 

Dicrocoeliidce,  morphology,  232,  265 
Dicrocceliuni  dendriticuni,  intermediate  host 

unknown,   267 

morphology,  266 

organs  of,  diagram  showing,  265 

ova  and  miracidia,  266 

synonyms,  266 

—  lanceolatum,  incidental  human  parasite,  7 
Diesing,  first  record  of  case  of  Metastrongy- 

lus  apri  in  man,  433 
Difdmus  tunensis,  cause  of  diarrhoea,  57 

characters,  57 

Difflugia  enchelys,  47 

Digenea,  morphology,  230 

Digestive  system,  loss  of,  in  parasites,  3 

Dinoflagellata,   52 

Dioctophyme,  431 

—  gigas,  hosts  and  habitat  in  body,  431 
— ■  —  in  man,   source  of,  431 

infection  by,  431 

morphology,  431 

ova  of,  432 

synonyms,   431 

Dioctophymidoe,  431 

—  characters,  375 

Dionis    de    Carrieres,    liver-fluke    in    right 

hypochondriac   region,   244 
Diphyllobothrium     (Bothriocephalus)    ditre- 

mum,  discovery  of  progeny  of,  13 
Diplodiscus  subclavatus,  hosts  of,  6 
Diplogonoporus,   morphology,   316 
—  grandis,    egg   of,   298 

morphology,  316 

ventral  view  of  genitalia  of  left  side, 

317 

of  portion  of  strobila,  317 

Diptera,   bibliography,   612 

—  biting-mouthed    and    other    noxious    car- 

riers of  disease,  600 

—  characters,  531,  532 

—  digestive  tract  of,  inhabited  by  Herpe- 

tomonads,  102 

—  larvae  of,  in  man,  references  to,  599 
Dipylidiidoe,   309,   320 

Dipylidium,  morphology,  320 

—  synonyms,   320 

—  caninum,  320 

confused  with  Taenia  solium,  660 

cysticercoids  of,  322 

hosts   of,   322 

dogs  and  cats  infected  with,  through 

skin  parasites,  323 

embryo  of,  development,  321 

expulsion   of,  drugs  suitable  for,   660 

hosts  of,  6,  7,  322 

invasion  by,  effect  on  central  nervous 

system,   649 

morphology,   320 

oncosphere  of,  299 

oncospheres    of,    animals    selected    as 

hosts  for  development,  299 


852 


THE   ANIMAL    PARASITES    OF    MAN 


Dipylidium  caninum,  prevalence  of  infec- 
tion by,   in   children,   659,  660 

proglottids  of,  322 

proglottis    of,    central    portion,    321 

region  of  human  body  inhabited  by, 

660 

rostellum  of,  289 

Dirksen,  effect  on  anaemia  of  expulsion  of 
Tosnia  solium,  648 

Dirofilaria  immitis,   417 

—  magalhaesi,  morphology,  417 

—  morphology,  416 

—  repens,   417 
Discophora,  see  Hirudinea 
Disporea  in  Myxosporidia,  182,   184 
Distoma,  opercula  of  ova  of,  discovery,  12 

—  echinatum,  redia  of,  in  later  stage,  227 
Distomata,  morphology,  231 

—  cercarice,  753 

Distomum  ophthalmohium,  244 

Doeveren,  van,  on  transmission  of  intestinal 

worms,   11 
Doflein,   Coccidiomorpha,    129,    151 

—  Cnidosporidia,    129,    194 

—  Entamceba    kartulisi,    44 

—  Trypanosoma    equiperdum,    97 
Dog  and  cat,   parasites  common  to,   6 

—  blood    of,    life-cycle    of    Babesia    (Piro- 

plasma)   canis  in,   175 

—  DipylidiuTYi  canimim  parasite  in,  322 

—  fleas,  natural  flagellates  in,   111,   112 
transmission   of   canine   kala-azar  by, 

103 

see     also     Ctenocephahis     canis     and 

Pulex  serraticeps 

—  host  of  Dibothriocephalus  latus,  313,  315 
of  Paragonimus  kellicotii,  250 

—  intestine  of,  Isospora  bigemina  parasitic 

in,  149 

—  mange,  transmission  to  man,  523 

—  rearing  of  Tcenia  echinococcns  in,  356 

—  to    dog,    transmission     of    leucocytogre- 

garine  from,  by  tick,  155 

—  transmission  of  infection   with  Demodex 

folliculorum  canis  to  man,   709 
Dogs,  contraction  of  surra  by,  9G 

—  dermal   infection   with   larvae  of  Ancylo- 

stomum  duodenale,   455 

—  echinococci  in,  346 

—  experiments  with,  to  prove  transmission 

of  infantile  kala-azar  by  fleas.   111 

—  infected       with       Dipylidium       caninum 

through  skin  parasites,  323 

—  infection     with    infantile     leishmaniasis, 

110 

experimental,  110 

natural,  110 

—  Leishmania  tropica  in,  108 

—  malignant  jaundice  in,  carrier  of,  493 
cause  of,  177 

—  nagana  fatal  to,  94 

—  percentage   infected   with   Tcenia   echino- 

coccns   in    various    cities    and    countries 
(footnote),   345 

—  prevention  of  echinococcus  infection  by, 

346 


Dogs,  segregation  of,  as  preventive  against 
Oriental  sore,  628 

—  skin    diseases    in,    due    to    young    nema- 

todes, 378 
Doliocystidce,    135 
Domestic     animals,     species     of     Sarcoptes 

transmissible  from,  to  man,  520 
trypanosomes  deleterious  or  lethal  to, 

69,   70 
Dormouse,      inoculation     of      Trypanosoma 

lewisi  into,  90 
Dornbliith,  method  of  evacuation  of  Oxyu- 

ridoe,  697 
Dorr,  tapeworm  drug  recommended  by,  674 
Dourine,  anaemia  and  paralysis  in,  97 

—  periods  or  stages  of,  97 

—  trypanosome   causing,   97 
Dovecots,  habitat  of  Argas  reflexus,  506 
Dracontiasis,  disorder  named  by  Galen,  386 
Draciinculidce,   385 

—  characters,    374 

Dracunculus     medinensis     (Guinea     worm), 
anterior   extremity,   387 

expulsion  by  extraction,  676 

—  methods  and  drugs  for,  676 

female,  transverse  section  of,  388 

—  —  infection  by,  prophylaxis  against,  676 
—  symptoms    and    lesions    following, 

389 

intermediate  host,  388 

larvae  of,  388 

life-history,  386,   388 

methods  of  extraction  from  body,  389 

morphology,  386 

period  of  development  in  man,  389 

regions  of  body  inhabited  by,  386 

synonyms,  386 

viviparous   nematode,   371 

—  morphology,  385 
Dromedaries,  "  mbori  "  in,  96 
Drone  fly,  583,  584 

Drosophila  melanogaster,  larvae  of,  charac- 
ters   and   habitat,   584 

effects    produced    by    ingestion    of, 

584 
Drosophilidoe,  characters  and  habitat,  584 
Drouillard,    favourable   effects   of  expulsion 

of  Ascaridoe,  649 
Drugs,  reaction  of  spirochsetes  to,  115 
Dubini,  Filaria  (?)  conjunctivae  in  man,  405 
Dubreuilh,    infection    with    Demodex    folti- 

culorum,  708 
Ducks,  destruction  of  mosquito  larvae  by,  636 

—  how  infected  by  Echinostoma  echinatum, 

226 
Dufour,    creation    of    name    Gregarina    by, 

129 
Duguet,    maculae    caeruleae     (tdches    bleues) 

due  to  infection  by  crab  louse,  712 
Dujardin,   development  of  Taeniae,  14 

—  psorosperms,   181 

Duke,   anterior   station   of  trypanosomes   in 
Glossince,   101 

—  Trypanosoma  gambiense  in   antelope,   76 
Dum-dum  fever,  105 


INDEX 


853 


Dumesnil,  larvae  of  Muscidce  in  nose,  720 
Duodenum,   flagellate  stage  of  Larnblia  in- 

testinalis   found  in,   59 
— -  human,     habitat    of    Ancylostoma    duo- 

denale,  450 

—  possible  invasion  by  Balantidium  7ninu- 

tuvi,    204 

—  species    of    Trichostrongylus    inhabiting, 

435,  436 
Durham,  specimens  of  Leptus  (bete  rouge), 

486 
Dutton,  Trypanosoma  gamhiense,  68 

—  and  Todd,  herpetomonads  in  mice,  738 
researches  on  Spirochoeta  duttoni,  116, 

117 
Duval,  liver-flukes  in  veins,  243 
Dysentery,   amoeba   as   causal   agent,   30,   34 
—  experiments  made  to  prove,  30, 

40,   618 

—  amoebic,  acute,  symptoms,  618 
carriers,   618 

chronic,  618 

experimental  production,  618 

gangrenous,  treatment,  619 

—  —  incubation  period  long,  618 
latent,  618 

leading  to  liver  abscess,  618 

preventive  measures,  620 

relief  of  griping  and  straining,  618, 

619 

—  ■ —  treatment  by  emetine   hydrochloride, 

618,  619 

by  ipecacuanha,  619 

by    preliminary    administration    of 

castor  oil,  619 
of  liver  abscess,  620 

—  —  —  surgical,   619 

—  association  of  Lamhlia  intestinalis  with, 

59,  625 

of  Noc's  entamoeba  vd^ith,  41 

of   Trichomonas   with,   56,   624 

—  bacillary,  31 

Entamoeba  coli  present  in  cases  of,  38 

—  bacillus,  discovery  of,  31 

—  balantidian  or  ciliate,   201,   202,   637 
prophylaxis,    637 

■  —  symptoms,  637 

treatment,  637 

—  discoveries      of      Cholera      Investigation 

Commission   with  regard  to,   30 

—  discovery  of  intestinal   amoebae  in  cases 

of,  30 

—  flagellate,  623 
diet  in,  625 

geographical  distribution,   624 

prognosis,    625 

prophylaxis,  625 

protection    of    food    supply     against 

fsecal  contamination  in,  625 
treatment,    624,    625 

—  followed  by  recovery  from  oxyuriasis,  698 

—  production  by  injection  of  amoebae  into 

cats,  35 

—  red,    geographical   distribution,    147,   741 
in  cattle,  cause  of,  147 

—  so-called,  due  to  mites,  512 


Dysentery  spread  by  house-fly,  586 

—  treatment  by  bismuth  subnitrate,  619 

Dysodius   lunatus,  bite  of,  541 


Ear,  abscess  of,  liver-fluke  in,  244 

—  parasites  in,  615 

see  also  Myiasis,  auricular 

Ears  of  hosts  infested  by  Ornithodorus  m6g- 

nini  rendered  painful,  510 
Earth-eating   in   connection   with   Trichuris 

trichiura  infection,  679 
East  Coast  fever  in  cattle,  cause  of,  178, 179 

pathogenic   agent,   174 

Eau  de  Cologne,  application  in  nasal  myia- 
sis, 719 
Echidnophaga,  543 
Echinococcus,  brood  capsules,  349 

and    scolices,    mode    of    formation, 

348,  349 

transformation   into  daughter  cysts, 

351,  353 

—  cysticus  jertilis,  350 

—  cysts  causing  urticaria,  651 

—  daughter  cysts,  350 

—  definition  of,  301 

—  fluid,  chemistry  of,  353 

—  frequency    of    infection    by,    of    various 

organs  in  slaughtered  animals,  347 

—  hominis  in  liver,  incised  fibrous  capsule 

and  wall  showing  daughter  cysts,  351 

—  in  dogs,  oxen,  sheep  and  pigs,  346 

—  in  man,   age   incidence,   355 

death   at   various    stages   of   develop- 
ment, 356 

—  —  geographical  distribution,  354,  355 

organs  of  body  invaded  by,  355 

percentage   of   prevalence    in    Central 

Europe,  354 

secondary,  356 

sex  incidence,  355 

—  infection,  prevention  of  spread  by  dogs, 

346 

—  m,ultilocularis  (alveolar  colloid),  develop- 

ment, 357,  358 
feeding  experiments  with  Taenia 

from,  358 

booklets  of,  359 

in  liver  of  ox,  357 

in    man,    early    disintegration, 

357 
geographical        distribution, 

358 

invasion,  results  of,  358 

_  _  _  site  of,  358 

morphology  of,  356 

reasons     for     distinction     from 

hydatid  or  unilocular,  357,  358 

—  of  liver  rupturing  into  abdominal  cavity, 

652 

—  rate  of  growth,  354 

—  rich  in  glycogen,  348 

—  scolex,  349 

development  in  rabbits,  353 


854 


THE   ANIMAL    PARASITES    OF    MAN 


Echinococcus  scolex  iu  jirocess  of  vesicular 
metamorphosis,  section  through,  351 

invaginated,   section   through,   350 

transformation     into     daughter     cyst, 

352 

—  scolices  in,  303,  349 

—  serum  diagnosis  of,  359 

—  structure  and  development,  347 

—  veterinorum,   350 

brood  capsules   and  scolices,  350 

hooklets   of,   355 

Echinorhynchus,  anatomy,  475 

—  excretory  organs,  475,  476 

—  "floating  ovaries"   of,   150 

—  gigas,  hosts  of,  477 

incidental  human  parasite,  7 

intermediate  hosts,  477 

morphology,  477 

—  hominis,  478 

—  moniliformis ,    hosts,    habitat    and    inter- 

mediate host  of,  478 

—  nervous  system,  475 

—  ova  of,  477 

—  protrusor  proboscidis,  475 

—  receptaculum  proboscidis,  475 

—  retractor  proboscidis,  475 
receptaculi,   475 

—  sexual   organs,   476 
Echinostoma,  cercarise  of,  225 

—  echinatum,  method  of  infection  of  duck.3 

and   geese   by,  226 

—  ilocanum,  habitat,  268 
morphology,   267 

organs  of,  diagram  showing,  268 

—  m,alayanum,  habitat,  269 
morphology,   268,   269 

—  morphology,  267 
EchinostomidxB,  267 

—  morphology,   233 
EchinostominoB,   267 

—  morphology,  233 
Ectoparasites,  1 

—  permanent,  changes  in,  3 
Ectoplasm  of  protozoa,  25,  26 

substances  deposited  in,  26 

Ectoschiza,  135 

Eczema  due  to  head  louse  infection,  709,  710 

—  following  infection  by  crab  louse,  712 

—  occupational,  diagnosis  from  scabies,  706 

—  peri-anal  and  perineal,  set  up  by  migra- 

tions of  Oxyuris  vermicularis,  695 

—  purulent,  following  infection  by  ancylo- 

stomes,  684 

—  resulting  from  clothes  louse  infection,  711 

—  set    up    by    infection    with    Dermanyssus 

gallince,   703 

by  Leptus  autiunnalis ,  702 

Ehrenberg,  Spirochceta  plicafilis,   114 
Ehrlich's  acid  hsematoxylin,  751 
Eimer,   researches  on   coccidia,    136 
Eimeria,  142 

—  avium,   causal   agent  of  white   diarrhoea 

or  white  scour  in  fowls,  and  blaclfhead 
in   turkeys,    145 

cause  of  fatal  epizootics  among  game 

birds  and  poultry,   142 


Eimeria   avium,  infection   by,  method  of,  145 

life-cycle   of,    142-145 

period,  144,  145 

phases,   142-144 

merozoites   of,   143 

microgametes    and    macrogametes    of, 

143,  144 

—  —  relation  to  E.  stiedoi,   145 

sporozoites  of,   143 

trophozoites  of,  143 

—  falciformis,   136 

—  (Coccidium)  schuhergi,  life-cycle  of,  138- 

141 

—  hominis,   150 

bodies  described  as,  150 

—  stiedce,    ascribed    cause    of    "  red    dysen- 

tery "   in   cattle,   147 

host    of,    7,    145 

oocysts    of,    142,     146 

parasitic    in    rabbit    and    occasionally 

in  man,   145,  148 

schizogony,    147 

—  —     effects,   147 

synonyms,   145 

see  also  Coccidiosis 

—  synonyms,   142 
Eimeridea,    141,    742 

Electrolytic  needle,  application  in  creeping 

disease,  731 
Elephantiasis    arabum    from    Filaria     han- 

crofti  infection,   sites   of  body   affected 

by,  677 

symptoms,  677 

treatment,   677 

—  scroti  in  filariasis,  402 

Ellermann,      "  rhizopods      in      poliomyelitis 

acuta,"  46 
Elmassian,  discovery  of  Entamoeba  minuta, 

42 
of       trypanosomes       in        "  mal       de 

caderas,"    68 
Embryophore  of  tapeworms,  298 
Emetine    and    vaccine    treatment    combined 

iu  pyorrhoea  alveolaris,  620 

—  hydrochloride  in  flagellate  dysentery,  625 

—  —  in    treatment    of    amoebic    dysentery, 

618,   619 

—  in  oral  endamoebiasis,  620 
Emily,  expulsion  of  guinea  worm,  676 

—  method    of    extraction    of    Dracnnculus 

medinensis ,  389 
Endamoeba,    31,    34,    734 

—  see  also  Entamoeba 
Endamoebiasis,   oral,   620 
treatment,  620 

Endermol,  application  in  scabies,  707 
Endoparasites,   1 

—  intermediate  generations  invading  inter- 

mediate  hosts,  5 
hosts   of,   5 

—  young  of,  leaving  host  or  organ  of  host 

inhabited  by  parents,  5 
Endophlebitis  set  up  by  Schistosoma  hcema- 

tobium,  274,  275 
Endoplasm   of  protozoa,   25,  26 
substances  deposited  in,  26 


INDEX 


855 


Endoschiza,   135 

Endotoxins  in  Trypanosoma  equiperdiim,  98 

Endotrypanurn  schaudinni,  99 

Enemata,  in  evacuation  of  Oxyuridce,  697 

Entamreba,  31,  734 

—  africana,  see  under  Entamoeba  tetragena 

—  bticcalis,  43,   734 

association  with  cancer  of  oral  cavity, 

43 

with  dental  caries,  43 

— with    pyorrhoea   alveolaris,   43,    734 

characters,  43 

possible    relation    of    E.     pulmonalis 

to,  45 

—  biitschlii,  34 

—  coli,    32,    618,    733 
characters,    33 

—  —  cysts   of,   33 

in  normal  faeces,  33 

encystment  process,  33 

— cytological    changes   during,    33 

how  distinguished  from  E.  histolytica, 

34,    40,    733 

life-cycle    of,    32,    33 

non-pathogenic     and     non-culturable, 

618 

parasite   of  human   intestine,  618 

may  be  present  in  bacillary  dysen- 
tery,   38 

schizogony    of,    32,    33 

so-called  autogamy  of,  34 

—  —  sporogony  of,   32,  33 

—  gingivalis,    733 
synonyms,    734 

—  hartmanni,  34 

—  histolytica,   32,   34,   45,   618,   733 

causal  agent  of  amoebic  dysentery,  35 

changes  in  intestine  produced  by,  35 

—  — ■  characters,  34,  35 

cysts  of,  permanent,  injection  pro- 
ducing infection,  37 

dysentery  following  experimental  in- 
fection with,  618 

how   distinguished    from   E.    coli,    34, 

40,  733 

present  in  large  intestine,  38 

—  —  producing  liver  abscess,  35,  620 
sporulation  of   (so-called),  37 

and   E.    coli,   differences   between,   34, 

40,  733 

mixed  infection,  38 

and    E.    tetragena,    identity    of,    38, 

40,   41 

—  hominis,  42 

—  kartulisi,  44,  734 

association  with  dental  caries,  44 

—  maxillaris ,  734 

—  minuta,  42 

relation  to  E.   histolytica,  40,  42 

—  mortinatalium,   45 

—  nipponica,  42 

—  Noc's,  41 

association     with     liver     abscess     and 

dysentery,  41 

—  phagocytoides,  42 

—  poleki,  34 


Entamoeba  pulmonalis,   45 

relation  with  E.   buccalis,  45 

—  tetragena,  38 
characters   of,   39 

described    as    E.    africana    by    Hart- 

mann,  38 

found  in  amoebic  dysentery,  38 

infection   by,  40 

multiplication  of,  39 

nucleus  of,  39 

'  part  of  life-cycle  of  E.  histolytica,  38 

reproduction  of,  39 

trophozoites,  39,  40 

and  E.  histolytica  identical,  38,  40,  41 

—  tropicalis,  41 

—  undulans,   43,    44 
characters  of,  43 

probable  flagellate  nature  of,  44 

—  williamsi,  34 
Entamoebse  of  vertebrates,  34 

—  question     of     cultivation     on     artificial 

media,  42,  742 
Enteritis,    hsemorrhagic,    in    Strongyloides 
stercoralis  infection,  674 

—  produced    by    injection    of    amoebae    into 

cats,  36,  37 

—  verminosa  in   children,  688 
Entozoa,  1 

—  derivation  of,  21 
Enyaliopsis   durandi,   bite   of,   542 

—  petersi,  542 

—  species  of,  producing  ulcers,  542 
Eosinophilia  in  ancylostomiasis,  647 

—  in  bilharziasis,  641 

—  in   hydatid  disease,  652 
Epicarin,  application  in  scabies,  707 
Epidermis,    human,    excavation    of    tunnels 

in,  by  Sarcoptes  scabiei,  var.   hominis, 
519 

—  "  wormlet "  burrowing  into,  599 
Epistaxis,  association  with  presence  of  Lin- 

guatula  in  nasal  cavity,  526,  527 

—  leeches  in  nose  causing,  701 
Epithelium  of  nematodes,  360 
Epizoa,  1 

EproboscidcB,  see  Pupipara 
Epstein,       experimental       infection       with 
Ascaris  lumbricoides ,  465 

—  transmission    of    trichomonad    infection, 

56 
Epstein's  method  of  diagnosis  of  ascariasis, 

692 
Equines,  baleri  in,  causal  agent,  95 

—  biliary  fever  in,  cause  of,  177 
Erdmann,      experimental      infection      with 

Sarcosporidia,    192 
Erythema,     autumn,      set     up     by     Leptut 
autum^nalis,  485,  486 

—  following  bite  of  Argas  reflexus,  506 

—  set    up    by    infection    with    Dermanyssus 

gallince,  703 

by  Leptus  autumnalis ,  702 

i    Eschatocephalus,  characters  of,  497 
I    —  hosts  and  habitat  of,  497 

Escomel,    dysentery    due    to    Trichomonas, 
I  56,  624 


856 


THE   ANIMAL    PARASITES    OF   MAN 


Escomel,  treatment  of  esjjundia,  629 

of    lamblial    diarrhoea,    625 

Espundia,    108,    628 

—  course  of,  629 

—  geographical  distribution,  628 

—  pathology  of,  628 

—  prophylactic  measures  against,  629 

—  transmission  of,   629 

—  treatment,   629 

Ether,  application  in  nasal  myiasis,  719 

—  lotions  in  crab  louse  infection,  712 

—  sulphuric,  in  ^  crab  louse  infection,  712 
Ethyl    chloride,    freezing    by,    in    creeping 

disease,  731 
Eucalyptus  oil  in  expulsion  of  ancylostomes, 

687 
Euflagellata,  52 

—  characters   of,   52 

Eugregarinea,  schizogony  absent  in,  134 
EuLyes  amoena,  542 

Euphorbia,      infection      by      Herpetomonas 

davidi,  104 
Eupodidoe,  characters,  491 
Euquinine  in  malaria,  635 
Europe,   Central,   percentage   of   prevalence 

of  echinococcus  in  man  in,  354 
Eustrongylus    gigas,   infection   by,   site   of, 

681 

symptoms,  682 

Evans,  discovery  of  trypanosomes  in  blood 

of   horses    with    "  surra "    disease,    67 

—  see    also    Steel   and   Evans 
Excretory  apparatus  of  cestodes,  291 

—  or  segmental  organs  of  Hirudinea,  481 

—  organs  of  Echinorhynchus,  475 

of  nematodes,  366,  367 

Eye,  cysticerci  in,  335,  664 

—  cysticercus  of,  664 

diagnosis  from  foreign  body,  664 

—  diseases  due  to  Liicilia  Tnacellaria,  721 

—  human,  infection  with  filaria,  406 

—  —  nematodes  observed  in,  412 

—  invasion  by  Loa  loa,  678 

—  paragonimiasis  of,  639 

Eyeball,  bodies  found  in,  probably  young 
liver-flukes,  244 

Eyebrows  and  eyelashes,  pediculosis  of, 
treatment,  712 

Eyelid,  iipper,  extraction  of  larva  of  Hypo- 
derma  hovis   from,   596 

Eyelids,  Demodex  folliculorum  affecting,  708 

Eyes,  destruction  of,  by  Sarcophaga  wohl- 
fahrti,   723 

—  loss  of,  in   parasites,  3 


Fabre,  prophylaxis  against  ancylostomiasis, 

685 
Face,  swelling  of,  in  nasal  myiasis,  718 
Faeces,   amoebse   found   in,   30,   34,   47,  48 

—  asexual  multiplication  of  Chlamydophrys 

enchelys  in,  47 

—  examination  for  protozoa,  746 

—  human,    fresh,    Strongyloides    stercoralis 

larva  from,  382 


Faeces,    larvae    of    Uomalomyia    canicularis 

found  in,  584 
of   Piophila    casei    found    in,    583 

—  males    of     Oxyuris    vermicularis    rarely 

met  with  in,   468 

—  normal,  cysts  of  Enta1no^ba  coli  present 

in,  33 

—  preservation  of  ova  of  flukes  in,  472 

—  N^orticella  in,  206 

Fanapapea       intestinalis       identical       with 

Tetramitus   mesnili,  57 
Fantham,  H.  B.,  appendix  on  protozoology 
(recent  researches,    formulae   of   culture 
media,     general     protozoological     tech- 
nique), 733 

avian    coccidiosis,    145 

pathogenic  spirochaetes,  119 

classification    of    Haplosporidia,    195 

of    Schizogregarines,    135 

experimental     infection     with     Spiro- 

chceta  duttoni,  117 

granule  phase  of  spirochaetes,  120 

Herpetomonas    ctenocephali,    103 

pedictili,    103 

latent  forms  of  trypanosomes,  73,  74, 

77 

molluscan    spirochaetes    breaking    up 

into  granules,  119 

—  —  morphology  and  life-cycle  of  Eimeria 

avium,   142-145 
and      life-history      of      Spirochoeta 

hronchialis,  739 
nuclear  phenomena  of  Babesia  hovis, 

176 

Protozoa,  25 

recent  work  on  spirochaetes  of  human 

mouth,  740,  741 

Rhinos poridium   kinealyi,    195-197 

schizogony    in    Leucocytozoon    lovati, 

153 
significance    of    insect    flagellates    in 

relation  to  disease,  104,  739 
Theileria  parva,  179 

—  —  Trypanosoma  rhodesiense,  69,  76 

■ and  Porter,  experimental  introduction 

of    insect    flagellates    into    vertebrates, 
104,   112,  738 

inoculation  experiments  with  Her- 
petomonas jaciilum-,  104,  738 

— natural     herpetomonads    in    mice, 

739 

researches  on  Nosema  apis,  185 

■ — ^  on  Spirochceta.  duttoni,   116 

'    —  —  and    Thomson,    J.    G.,    cultivation    of 
Babesia  canis,   172 

periodic    cyclical    variation    of 

trypanosomes   in   blood,   78 

see  also  Stephens  and  Fantham 

Fasciola,   237 

—  gigantica,  distribution,  244 
habitat,    244 

invading  lung,   245 

morphology,   244 

—  section  illustrating,  243 

synonyms,  244 

—  hepatica,  237,   238,  239,  638 


INDEX 


857 


Fasciola   hepatica,  cercaria  of,   228 

encysted,   228 

development   of,   226 

fixtition  method,  471 

geographical  distribution,  238 

half  transverse  section  through,  214 

hosts  of,  6,  7,  238 

incidental  human  parasite,  7 

intermediate  host,  240,  241 

invading  and  infecting  pharynx,   242 

regions   of  body  other  than   liver, 

243 
life-history,  241 

—  —  method  of  infection  of  sheep  by,  226 
miracidium  of,  223 

morphology,   237 

sections  illustrating,  238,  239 

ova  of,  to  be  distinguished  from  those 

of  Dibothriocephalus  latus,  242 

ovum  of,  223 

from  liver  of  sheep,  240 

redia,   in   early   stage,   227 

—  —  synonyms,   237 

—  morphology,  237 

Fascioliasis,  prevention  and  treatment,  G38 

—  symptoms  of,  638 

—  see  also  Liver-fluke  disease 
Fasciolidce,  237 

—  morphology.  231 
Fasciolince,  237 

—  morphology,  231 
FasciolopsincB,  245 

—  morphology,   231 

Fasciolopsis  buski,  fixation  method,  471 

geographical  distribution,  246 

habitat,   246,   638 

morphology,  245,  246 

symptoms  set  up  by  invasion  by,  638 

—  fiillehorni,  cirrus  sac,  247 

habitat,  249  ' 

morphology,  247 

—  ventral   aspect  showing,   248 

—  goddardi,   morphology   and   geographical 

distribution,  247 

—  rathouisi,  geographical  distribution,  247 
habitat,   247 

morphology,  246 

—  morphology,  245 
Feltinella,  characters,  561,  567 
Females,   greater  prevalence  of  head  louse 

infection  among,  709,  710 

—  rarity  of  bilharziasis  in,  643 

Fern  root,  new  species,  effects  as  vermi- 
fuge, 673 

Fibrin,  clots  of,  Dioctophyme  gigas  in  man 
traced  to,  431 

Ficalbia,    characters,    565 

Fievre  de  grain,  702 

Fiji,   intermediate  host  of  filaria  in,  575 

—  manifestations  of  filariasis  in,  402 
Filaria  associated  with  phthisis,  408 

—  hancrofti,    anatomy   of,    diagrams   show- 

ing, 391 

discoveries  relating  to,   390 

diseases   following   infection   by,   398i 

400 


Filaria  hancrofti,  embryos,  392 

female,  characters,  392 

geographical  distribution,  392,  403 

habitat  In  body,  392 

■ infection  by,  diseases  resulting  from, 

676,  677 

larvae  of,  absence  from  blood  in  those 

suffering  from  filarial  disease,  400 

distribution   in  body,  392,  393 

method  of  concentration,  395 

of  preservation,  395 

morphology,  determination  by  fixa- 
tion and  staining  methods,  395,  396 

— periodicity  of,  in  peripheral  blood, 

393,  394 

separation  of  red  corpuscles  from. 


of    Tseniorhynchus    carry- 


395 

species 

ing,  577 

—  —  —  structure,  396 

life-history,  398 

male,  characters,  392 

mosquitoes  acting  as  hosts  of,  398 

ova  of,  392 

synonyms,  390 

transmission  of,  576 

—  (?)  conjunctiva,  404,  405 

—  —  morphology,  404 
normal  hosts  of,  406 

sites  of  infection  in  man,  405 

synonyms,  404 

—  demarquayi,  403,  404 

geographical  distribution,  403 

morphology,  403 

—  infection  of  skin  by,  378 

—  intermediate  host  of,  in  Fiji,  575 

—  (?)  kilimarce,  407 

—  loa,  host  of,  601 

—  medinensis,  antiquity  of  knowledge  con- 

cerning, 386 

—  morphology,  390 

- —  oculi  humani,  association  with  cataract, 
406 

—  perstans,  carrier  of,  508 

—  (  ?)  romanorum  orientalis,  morphology,  407 

—  (?)  sp.   (?),  407 

—  taniguchi,  404 

Filariasis,  cultivation  of  bacillus  from  cases 
of,  755 

—  prevalence  proportionate  to  prevalence  of 

Mikrofilaria  hancrofti  in  blood,  400 
Filariidce,  390 

—  characters,   374 
Filariinoe,  390 

Filmaron,  administration  in  expulsion  of 
ancylostomes,  687 

—  as  vermicide,  672 

—  oil,   dosage  of,   672 
effects  of,  672 

in  expulsion  of  Ascarides,  694 

''Filterable    viruses,"    207 

Filtration  experiments  with  CMamydozoon 
granules,  209 

Finlaya,  characters,  564 

Finsen,  echinococcus  cysts  causing  urti- 
caria,  651 


858 


THE   ANIMAL   PARASITES    OF   MAN 


Finucane,  lymphangitic  symptoms  ia  chil- 
dren from  Filaria  hancrofti  infection, 
676 

Fischer,  effects  of  new  species  of  fern  root 
as  a  vermifuge,  672,  673 

—  retinal      haemorrhages      in      ancyloatome 

anaemia,  646 
Fish,  destruction  of  mosquito  larvae  by,  636 

—  disease    of,    due    to    invasion    by    Myxo- 

sporidia,  182,  184 

—  eating  of  raw  or  badly   cooked,   favours 

transmission  of  Dibotliriocephalus  latus 
in  man,  315 

—  fresh-water,  second  intermediate  host  of 

Clonorchis  endemicus,  261 
proved   by  feeding  ex- 
periments, 261 

—  intermediate    host    of   Dihothriocephalus 

latus,  255 
of  Opisthorchis  felineus,  255 

—  parasitic  ciliate  destructive  to,  206 

—  plerocercoids  of  Dibothriocephalus  latus 

inhabiting,  314 

—  trypanoplasms  in,  68 

Fistulas,  anal  and  rectal,  set  up  by  migra- 
tion of  Oxyuris  vermicularis,  695 

—  formation  of,  through  migration  of  para- 

sites, 9 

—  urethral,  arising  from  bilharziasis,  642 
■ — treatment,  644 

Fixation,  time  of,  749 
Fixatives,   748,   749 

—  for  wet  films,  748 

—  hot,  748 

Flagella  may  occur  among  rhizopods,  52 

—  of  Flagellata,  50,  51 
Flagellata,  50 

—  characters  of,  28,  50,   51 

—  classification,  52 

—  formation  of  colonies  of  individuals,  51 

—  habitat,   28,  52 

—  multiplication  of,  51 

—  non-flagellate  stages,  51 

—  nuclear  apparatus  of,  51 

—  post-flagellate   and  pre-flagellate   stages, 

52 
Flagellates,  aggregation  rosettes  of,  51 

—  dysentery  in   children   due  to,   56,   624 

—  in    blood    of   horses,    diseases    associated 

with,  67,  68 

—  natural,  in  dog  fleas,  112 

of  invertebrates,  evolution  of  Leish- 
lYiania  from,  739 

—^  parasitic,  in  insects,  experimental  intro- 
duction into  vertebrates,  104,  112,  737, 
738 

relation    to    evolution    of 

leishmaniasis,  112,  737,  738 

in  relation  to  evolution  of  disease,  739 

Flagellosis  of  plants,  possible  connection 
with  leishmaniasis,   104,  739 

Flat  worms,  see  Platyhelminthes 

Flea,  human,  see  Pulcx  irritans 

Fleas  acting  as  intermediate   hosts,   543 

—  blood-suckers,  543 

—  carriers  of  plague,  543 


]'^leas,  cocoons  of,  543 

—  fed  on  infected  rat,  percentage  infected 

with  trypanosomes,  93 
--  herpetomonads  in  gut  of,  103 

—  larvae  of,   543 

—  life-cycle  of  Trypanosoma   lewisi   in,  88,  90 
— ■  method    of    controlling,    during    experi- 
ments,  93 

—  ova  of,  543 

—  jjossible  transmission  of  kala-azar  by.  111 
Flemming's  solution,  749 

Flesh  of  animals  containing  larvae  of  tape- 
worms must  be  thoroughly  cooked  be- 
fore eating,  668 
Flesh-fly,   see  Sarcophaga   carnosa 
Flexner,  Entamoeba  kartulisi,  44 
Flies,   larvae  of  different  species  of,   found 

in   intestinal  myiasis,  728 
Flukes,  clearing  and  mounting  agents,  471 

—  differentiation  methods,  471 

—  fixation  methods,   471 

—  miracidia   of,    discovery,    12 

—  ova    of,    preservation    in    faeces,    urine, 

bile,  472 
transference  to  glycerine,  472 

—  preservation  and  examination  of,  471 

—  staining  methods,  471 

—  see  also  Trematoda 

Foetus,  Trypanosoma  cruzi  in,  88 

Foley,  transmission  of  relapsing  fever,  120, 
121 

Food,  transmission  of  trichomonad  infec- 
tion by,  56 

Foods,  decomposing,  inhabited  by  and 
nutriment  of  TyroglyphidcB,  511 

Foot,  sole  of,  hepatic  flukes  found  in  swell- 
ing  on,   243 

Foot-and-mouth    disease,    207,    208 

Foraminifera,  27,  47 

—  characters  and  habitat,  27 

Forde  and  Dutton,  discovery  of  human  try- 
panosomes, 68 

Foreign  body,  diagnosis  of  cysticercus  of 
eye   from,   664 

Formalin,  fixation  of  cestodes  by,  472 

Fowler,  method  of  administering  male  fern 
to  children,  671,  672 

Fowler's  solution  in  sleeping  sickness,  623 

Fowls,  fatal  effects  of  Spirochoeta  galli- 
narum  on,  119 

Fox,  host  of  Dibothriocephalus  latus,  313 

Framboesia   tropica,    see    Yaws 

Fran9a,  action  of  leucocytozoa  on  red  cells, 
153,   742 

—  genera  of  Piroplasmidce,   174 
Francaviglia,  auricular  myiasis,  615 
Franchini,   experimental  infection  of  verte- 
brates   with    herpetomonads,    103,    104, 
112,  739 

—  Hcemocystozoon    brasiliense,    104 
Frese,  O.,  rhabdites  found  in  gastric  fluid 

obtained  by  lavage,  378 

Freund,  sarcophaga  larvae  from  abscess 
cavities,  723 

Frog,  rectum  and  bladder  of,  ciliates  para- 
sitic  in,   207 


INDEX 


859 


Frontal    sinus,     invasion     by    Ancylostoma 

duodenale,  683 

scolopendra  in,  721 

Fruit  pickers  affected  by  Leptus    autumna- 

lis    (footnote),   485 
Fiilleborn,    cultivation    of    larval    forms    of 

Ancylostoma  and  Strongyloides,  474 
Furcocercous   cercarise,    753 
Fiirst,   cases  of  Ascarides   invading-  larynx 

and  trachea,  691 


Gabel,   diarrhoea  due   to   Difdmus   tunensis, 
624 

—  Difdmus   tunensis,  57,  624 
Gad-flies,  600 

—  see  also  TabanidcB 

Gaetano,  cysticercus  of  tongue,  663 
Galen,  disorder  named  "  dracontiasis  "  by, 

386 
Gall-bladder,       Schistosoma      hcematohium, 

eggs  of,  in,  274 
Gall  sickness  in  cattle,  cause  of,  98,  180,  611 
Galleria  melonella,  larvae  of,  in  nose,  720 
Galli-Vallerio,  bothriocephalus  anaemia,  646 

—  infection  by  trichomonads,  56 

—  Oxyuris  and  Trichocephalus  infection  in 

relation  to  appendicitis,  654 
Galyl  in  syphilis,  632 
— ■  in  trypanosomiasis,  622 
Gamasidce    (coleopterous    or    insect    mites), 

characters,  491 

hosts  and  prey  of,  491 

Gambia  horse  sickness,  cause  of,  100 
Game  infested  by  Dermacentor  reticulatus, 

503 
Game-birds,  fatal  epizootics  among,  due  to 

Eimeria   avium,    142 
Gametes  of  Coccidiidea,  137,   139,   140 

—  of  gregarines,   132,   133 

—  of    malarial    parasites,    162 
Gametocytes   of   Coccidia,    140 

—  of    gregarines,    132,    133 

—  of  malarial  parasites,   162 
Gametogony,  in  Eimeria  avium,  143 
Gam,marus   pulex   occasionally   parasitic   in 

man,   483 
Garlic    and    saline    injections    in    Trichuris 

trichiura  infection,  680 
GastrodisciidcB,  236 

—  morphology,  231 

Gastrodiscoides,     how     distinguished     from 

Gastrodiscus,  236 
Gastrodiscus,   236 

—  mgyptiacus,  hosts  of,  237 

—  ho  minis,  236 

genital  pore,  236 

geographical  distribution,  237 

■  habitat,  237 

morphology,   236 

ova,  237 

testes,  236 

—  male  and  female  genitalia,  236 

—  morphology,  236 
Gastrophilus,  599 


Gastrophilus  equi,  599 

—  hcemorrhoidalis ,  599 

—  larvae  of,  in  stomach  and  intestine,  599 

—  nasalis,  599 

—  pecorum,  599 

Geber,  treatment  of  crab  louse  infection,  712 
Gecko,  blood  and  organs  of,  herpetomonad 

flagellate  in  cultures  from,  739 
Gedoelst,  Cordylobia  rodhani,  593 
Geese,     how     infected     with     Echinostom,a 

echinatum,  226 
Genital   apparatus  of  Cestoda,  293-296 
Genitalia      as      means      of      distinguishing 

species  of  Glossina,  604 
Genitals,     external     female,     invaded     by 

Oxyuris  vermicularis,  467 
Genser,  von,  Ascaris  infection  in  relation  to 

appendicitis,  653 
Gentian  violet,  752 
Gerbillus  indicus,  154 
Gerlach,    stages    of    liver-fluke    disease    in 

sheep,  240 
Germany,  districts  of,  j3ercentage  of  occur- 
rence of  echinococcus  in  man  in,  354 

—  trichinosis  in,  epidemics  of,  423,  429 

—  —  prophylaxis  against,  429 

Giard,  microscopical  investigations  of  con- 
jugation in  gregarines,  130 

Giardia   (Lamblia)  intestinalis,  736 

Giemsa's   stain,   751 

formula  of,   751 

Giesker,  liver-fluke  in  sole  of  foot,  243 

Gilesia,  characters,  564 

Gingivitis,  nematode  larvae  in  periosteum 
of  upper  jaw  associated  with,  378 

—  see  also  Entamoeba  gingivalis,  733;  and 

E.   buccalis,  43 
Girard,  effects  of  Trichocephalus  infection, 
651 

—  Trichocephalus    infection    in    relation    to 

appendicitis,   653 
Glas,   cysticercus  of  tongue,  663 
Glossina   austenii,    characters,   605 

—  brevipalpis   group,   characters,   606,   607 

—  caliginea,  characters,  605 

—  characters,  603,  604 

—  development    of   trypanosomes    in,    101 

—  fusca,  characters,  606 

—  —  group,  characters,  606 

—  fuscipleuris,  characters,  606 

—  habitat  of   species,  605 
— ■  larvae  of,  604 

—  longipalpi.-i,  characters,  606 
head  of,  664 

—  longipennis ,  characters,  607 

—  medicorum,   characters,   607 

—  morsitans ,  characters,  606 

development    of    Trypanosoma    brucei 

in,  94 
development    of    Trypanosoma    rhode- 

siense   in,    percentage    of   fly   becoming 

infective,  82 
developmental   cycle  of   Trypanosoma 

rhodesiense  in,  81,  82 

—  —  geographical  distribution,  608 

group,  characters,  605 

method  of  reproduction,  604 


54 


86o 


HE    ANIMAL    PARASITES    OF    MAN 


Glossina    morsitans,   race  submorsitans,   609 
transmission     of      nagana      (tsetse-fly 

disease)    by,    93 
of     Trypanosoma    rhodesiense    by, 

69,  81,   608 

—  nigrofiisca,  characters,   606 
■ —  pallicera,  characters,  605 

—  pallidipes ,    antenna   of,   604 

—  —  characters,  606 

—  palpalis,    100 

blood-sucking,  607 

carrier  of  sleeping"  sickness,  605,   607 

—  —  characters,  605 

development     of     Trypanosoma     gam- 

biense  in,  74,  75 

geographical  distribution,  607 

group,  characters,  605 

larval  and  pupal  stages,  608 

—  —  proportion  becoming  infected,  608 
salivary  glands  of,  invasion  by   Try- 
panosoma gavibiense,  75 

transmission   of  sleeping   sickness   in- 
fection by,  68 

—  puparia  of,  604,  605 

—  special  means  of  distinguishing  species, 

604 

—  species  of,  artificial  infection  with  human 

trypanosome,    60.5 

—  spread  of  trypanosome  diseases  by,  603 

—  tabaniformis,  characters,  606 

—  tachinoides,  characters,  605 
Glycerine,  mounting  agent  for  flukes,  472 

—  transference  of  ova  of  flukes  to,  472 
Glycerophosphates  and  arsenic  in  bronchial 

spirochaetosis,  633 
Glychaemalum,  Mayer's,  751 
Glyciphagi,      differentiation      from      Tyro- 

glyphi,  513 

—  buski,  513 

—  cursor,   513 

—  domesticus ,  cause  of  grocer's  itch,  513 

—  hippopodes,   513 

—  prunorum,  513 

Glycogen,  echinococcus  rich  in,  348 

Gnathobdellida,  481 

Gnathostoma  hispidum,  hosts  of,  385 

—  morphology,   384 

—  siamense,    infection    by,    associated    with 

tumour  of  breast,  385 
morphology,  384 

—  sp.,  hosts  of,  385 

—  spinigeriim,   385 

hosts  of,  385 

morphology,   385 

Gnathostotnidce,  384 

—  characters,  374 
Gnats,  see  Culex 

Goebel,  bilharziasis,  641,  642 

Goeldia,  characters,  565 

Golden  beetle,  intermediate  host  of  Echino- 

rhynchiis  gigas,  478 
Goldmann,   male   fern  extract   in   expulsion 

of  Strongyloides  stercoralis ,  675 

—  sebirol  as  vermicide,  672 

—  taeniol  in   ancylostomiasis,  686 


Goldschmidt,  excretory  apparatus  oi  Ascaris 
lumbricoides ,  367 

—  formation  of  ova  of  trematodes,  223 
Golgi,  description  of  asexual  cycle  in  blood 

in  case  of  quartan  parasite,  157 
Gonder,    relation    of   infantile    kala-azar   to 
Oriental  sore,   108,   109 

—  strain     of     Trypanosoma     lewisi     losing 

resistance    to    arseno-phenyl-glycin,    03 

—  Theileria  parva,  178 
Gordiidce,  375 

—  characters  of,  479 

—  larvae  of,  479 
Gordius   aquations,  479 

—  chilensis,  479 

—  pustulosus,  479 

—  species  invading  man,  479 

—  tolosanus ,  479 

—  tricuspidatus ,  479 

—  varius,  479 

—  villoti,  479 

—  violaceus,  479 
Grabhamia,  characters,  564,  576 

—  do7'salis,   576 

—  geographical  distribution,  "576 

—  sollicitans,  geographical  distribution,  576 
Graffe,    escape    of   ascarides    from    inguinal 

tumour,  656 
Granate  root  as  vermifuge,  673 
Granuloma  inguinale,  spirochsete  associated 

with,  122 
Grass,  harvest  or  gooseberry  mite,  see  Lep- 

tus  autuynnalis 
Grassi,  Cercomonas  and  Trichomonas,  54 

—  development   of    cestodes    without   inter- 

mediate host,  17 
of  Trichuris  trichiura,  420 

—  discovery  of  amabse  in  stools,  30 

—  experimental  self-infection  with  Oxyuris 

vermicularis,  469 

—  expulsion  of  Hymenolepis  nana,  661 

—  Hymenolepis  nana,  324 

—  larval    stage    of   Hymenolepis    diminuta, 

327 

—  mosquitoes  in  relation  to  human  malaria, 

158 

—  on  Entamoeba  coli,  32,  33 

—  self-infection  with  Ascaris  lumbi'icoides, 

465 
Great  black  bug  of  Pampas,  see  Conorhinus 

renggeri 
Gregarina  blattarum,  135 

—  longa  from  larva  of  crane-fly,  130 

—  munieri,    from    Chrysomela    hcemoptera, 

131 

—  ovata,  host  of,  135 
Gregarines,  ectoplasm  of,   131 

—  endoplasm  of,  131 

—  gametes  of,   132,    133 

—  gametocytes  of,    132,   133 

—  mode   of   infection,    134 

—  monocystid,   130 
hosts  of,  130 

—  morphology  of,   130 

—  movements  of,  131 

—  myonemes  of,  130,  131 


INDEX 


8^1 


Gregarines,   polycystid,   130,   131 

protomerite,     deutomerite     and     epi- 

merite  of,  131 

—  resistant  spores  of,   purpose  of,   134. 

—  spore-production,  132,  133 

—  sporocyst  of,   134 

—  sporozoites  of,   132,   133 

—  syzygy   of,   132 

—  trophozoites  of,  132,  133 

—  zygotes  of,   132,   133 

Gregarinida,  characters  and  habitat,  28, 
130 

—  classification,    134 

—  history  of  discoveries  relating  to,  129 

Grey  ointment  in  crab  louse  infection,  712 

Grocer's  itch,  cause  of,  513 

Ground-itch,  skin  affection  set  up  by  in- 
vasion of  larvae  of  Ancylostomum  duo- 
denale,   455 

—  treatment,  754 
Guarnieri's  bodies,  207,  209 

Gubler,  case  of  human  hepatic  coccidiosis, 

148 
Guermonprez,     method     of     expulsion     of 

ascarides,  692 
Guinea-pigs,    experimental    infection    with 

Sarcocystis   muris,   192 

—  natural  occurrence  of  Paraplasma  in,  180 
Guinea  worm,  see  Dracunculus  medinensis 
Gurley  on  Myxosporidia,  182,  183 


H. 


H^MADIPSA,    482 

—  blood-sucking  pest  in  tropics,  482 
Haemagogus,  characters,  565 
Hsemalum,  Mayer's,  751 
Haemamoeba,    151,   742 
Haemaphysalis,  characters  of,  497 

—  leachi    (dog  tick),   carrier  of  malignant 

jaundice  in  dogs,  493 

transmitting  agent  of  Babesia  canis, 

177 

—  ^punctata,    characters    and    morphology, 

502 

—  —  hosts  of,  503 
synonyms,  502,  503 

Haematein,    essential    principle    of    haema- 
toxylin,   751 

—  solutions    of,    in   staining   flukes,    471 
Hoematobia  irritans,  610 
Hoematopinus  spinulosus,  88 

effect  on  strain  of  Trypanosoma  lewisi 

being  passed  through,  93 
Haematoxylin,   Delafield's    (or  Grenacher's), 

751 

—  Ehrlich's  acid,  751 
Haematuria  in  bilharziasis,  641 
Haementaria,   482 

—  officinalis  used  medicinally,  482 
Hmmocvstozoon  brasiHense,  104 
Haemoglobinuria,  infectious,  in  cattle,  cause 

of,  177 
Hmmogregarina  balfouri  (jaculi),  154 

—  gerbilii,    154 


Uoemogregarina  marceaui,   154 

—  nobrei,  154 

—  schaudinni,   var.    africana,   154 
Haemogregarines,   characters  of,   154,   742 

—  hosts  of,   153 

—  in  red  blood  corpuscles,  154 

—  leucocytic,  154 

—  transmission   of,   153 

—  variation  in  size  and  appearance  of,  154 
Haemolysis,      cure,       after      expulsion      of 

Ascaridce,  649 

Hcemonchus   contortus,   diseases  due  to  in- 
vasion by,  437,  438 

geographical  distribution,  437 

habitat  and  hosts  of,  437 

life-history,  438 

morphology,  436,  437 

symptoms    caused    by    invasion    mis 

taken  for  those  of  ancylostomiasis,  438 

— •  morphology,  436 

Hcemoproteus    (Halteridium)    columbce,    in- 
sects transmitting,    151,   612 

life-cycle  of,  152 

danilewskyi,   152 

Haemorrhoidal    veins.    Schistosoma    hoem,a- 
tobium  in,  273 

superior,  plexus  formed  by,  in  rectum, 

272 

Haemorrhoids     set     up     by     migrations     of 
Oxyuris  vermicularis ,  694 

H<Bm,osporidia,  151 

—  Babesia  or  Piroplasma  type,  154 

—  characters   and   habitat,   28,    151 

—  Haemogregarina  type,  153 

—  Halteridium  type,  151 

—  Leucocytozoon  type,   152 

—  Plasmodium  or  Haemamoeba  type,  151 

—  recent  views  regarding,  742 

—  transmission  by  Ixodince,  704 
Hagen-Thorn,  cysticerci  in  brain,  665 
Hair,  methods  of  getting  rid  of  nitg  from, 

710 

—  follicles,  mites  of,  522 

Hake,  discovery  of  Coccidia  by,  135 
Halipegus  ovocaudatus ,  host  of,  6 
Halteridium    parasites    occur    in    blood    of 

birds,  151 
"  Halzoun,"  affection  of  pharynx,  produced 

by  Fasciola  hepatica,  242 
Hampshire,  sand  flies  biting  in,  579 
Hanau,    Oxyuris    infection    in    relation    to 

appendicitis,    654 
Haplosporidia,   129,   194 

—  characters  and  habitat,   29,   194 

—  classification,   195 

—  life-cycle,    194 
Haplosporidium,    194,    195 

—  heterocirri,   195 

Harington,  guinea  worm  infection,  676 
Harlej,  treatment  of  bilharziasis,  643 
Harmostomum       leptostomum,       immature 

specimen,  215 
Harpactor  cruentas,  542 
Harris,    Penn,    liver-flukes    in    abscess    of 

occiput,  243 
Hartmann,  case  of  Oxyuridce  in  nose,  696 


862 


THE   ANIMAL    PARASITES    OF    MAN 


Hartmann,  indeiiendent  description  of 
Entamoeba  tetragena  under  name  of 
E.   africana,   by,   38  | 

—  multiplication  of   Trypanoso7na   cruzi  in 

vertebrate  host,  85 

—  toxic  action  of  Oxyuris,  651 

—  and  Chagas,   Cercomonas  parva,  737  i 

—  and  Whitmore,  formation  of  amoebulse,  34 
Hata,  modification  of  Noguchi  technique  for 

cultivation    of    spirochaetes    and    trepo- 
nemes,  126 
Hausmann,   effects   of  trichocephalus   infec- 
tion, 651 
Head  louse,  see  Pediculus   capitis 
Headache  in  nasal  myiasis,  717,  718 
Heart-water  fever  in  sheep,  carrier  of,  493 
Hectopsylla,  543 

Heekes,  Oxyuris  in  appendix,  655 
Heidenhain-Rosenbusch,     iron-hsematoxylin, 

752 
Heliozoa,  characters  and  habitat,  27 
Heller,  life-history  of  Oxyuris  vermicularis , 
467,  469 

—  percentage  of  rats  infected  with  Trichi- 

nella,  427 
Helmerisch's      ointment,      application      in 

scabies,  706 
Helminthes,  2 

—  biological,  not  systematic  group,   2,  3 

—  dead,  decomposition  of,  9 

—  growth  and  agglomeration  in  host,  9 

—  life  spent  in  intermediate  and  final  host, 

18 

—  life-history  of,   18 

—  multiplication  of,  discovery  of  method,  13 

—  origin  of,  discoveries  as  to,  10,  11 

—  ova  of,  mistaken  for  coccidia  (footnote), 

137 

—  permanent  parasites,  2 

—  producing  substances  toxic  to   host,   9 

—  separation  of  LinguatulidcB  from,  2 
Helminthiasis,  9 

—  meningitiformis,  649 

Helsingfors,  clinic  of,  work  done  on  bothrio- 

cephalus  anaemia  at,  644 
Hemiptera,  characters,  531,  532 

—  digestive  tract  of,  inhabited  by  Herpeto- 

monads,  102 

—  sub-orders  (footnote),  532 

Henle,  investigation  of  Gregarinida,  129 
Henneberg,  site  of  cysticerci  in  brain,  665 
Henoch,  expulsion  of  ascarides,  693 
Hepatozoon,   154 

Heptaphlebomyia,  characters,  565 
Herbst,     experimental    infection     with    en- 
cysted Trichinellffi,  423 
Hermann,    eucalyptus    oil    in    expulsion    of 

ancylostomes,  687 
Hermaphroditism  in  permanent  parasites,  4 
Herpetomonad     flagellate     in     cultures     of 

blood  and  organs  of  gecko,  739 
Herpetomonads,    experimental    infection    of 
birds  with,  739 

—  hosts  of,  102 

—  in  blood  of  birds,  739 

—  in  mice,  738,  739 


Herpetomonads,  stages  of,   103 
Herpetomonas,   67,    102 

—  ctenocephali,  103,  111,  112,  738 
inoculation   experiments  with,   103 

—  davidi  infecting  plant  genus  Euphorbia, 

104 

—  jaculum,    inoculation    experiments    with, 

104,   738 

—  life-history,    stages    of,    103 

—  muscoi  domesticce,   102,  739 

—  pattoni,  103,  739 

inoculation   experiments  with,   103 

—  pediculi,  103,  738 

—  species  of,  introduction  into  vertebrates, 

103,  104,  112,  738,  739 

—  stratiomyice,  738 
Herpetomoniases,    112,    738 

Hessler,  R.,  Norway  itch  (scabies  nor- 
vegica),  519,  520 

Heterogony  in  nematodes,  372 

Heterokaryota,   198 

Heterophyes  heterophyes ,  geographical  dis- 
tribution, 264 

habitat,  264 

morphology,  263 

organs  of,   diagram  showing,  263 

synonyms,   262 

—  morphology,  262 
Heterophyiidw,  262 

—  morphology,  232 
Heterotricha,  29,  200 

Uexactinomyxon  psammoryctis ,  spore  of,  187 
Hexamastix    ardin-delteili,   624,    735 
Hexapoda,  classification,  431 

Higueron,  milk  of,  in  ancylostomiasis,   754 
Hilton,  J.,  observation  of  encapsuled  Trichi- 

nellae,  423 
Himasthlince,  269 

—  morphology,  233 

Hindle,  avian  pathogenic  spirochaetes,  119 

—  experimental       infection       with      Spiro- 

chceta  duttoni,  117,  118 
Hippius     and     Lewinson,     relationship     of 

Oxyuridce  to  appendicitis,  698 
Hippobosca  camelina,  611 

—  capensis,   611 

—  equina,   611 

—  maculata,  611 
bite  of,  611 

—  rufipes,  98 

—  wings   of  various   species,   611 
HippobflscidcB,  611 
Hirudinca,  480 

—  alimentary  canal  of,  480 

—  anatomy  of,  480 

—  body  cavity  of,  480 

—  cocoons  of,  481 

—  excretory  or  segmental  organs  of,  481 

—  muscular  system  of,  480 

—  nervous  system  of,  481 

—  oesophagus  of,  480 

—  pharynx  of,  480 

—  sexual   organs   of,   481 

Hirudo  granulosa,  used  medicinally,  482 

—  medicinalis ,     geographical     distribution, 

481 


INDEX 


m 


Hirudo   medicinalis ,    habitat,    481 
morphology,  481 

—  morphology,   481 

—  niysomelas,   used  medicinally,  482 

—  troctina,  characters,  482 

geographical  distribution,  482 

Histiogaster,  characters,  515 

—  (entomophagus?)  spermaticus,  515 
—  habitat,  516 

synonyms,   516 

—  food   of,   515 
Histoplasma,   112 

—  association   with  splenomegaly,   112 

—  capsiilatum,   112,   739 

Hoffmann,  formation  of  spores  in  Treponema 

pallidum,   125 
Holostomata,  ova  of,  development,  224 
Holostomum  variahile,  hosts  of,  6 
Holothyrus  coccinella,  effects  of  bite  of,  493 
Holotricha,  29,  199 
Romalomyia  canicularis,  larvae  of,  584,  585 

characters  and  habitat,  584 

- —  scalaris,    585 

Homoptera   (footnote),   532 

Hoplopsyllus    anom,alus,    carrier    of    plague 

bacillus,  543,  547 

—  distinctive  characters,  545,  547 
Horse,  piroplasmosis  in,  cause  of,  174 

—  serum  in  cultivation  of  Treponema  palli- 

dum, 126 

—  sickness  (Gambia),  cause  of,  100 
Horses  attacked  by  Leptus  autumnalis,  486 

—  nagana  fatal  to,  94 

—  organs   infected   with   echinococcus,    per- 

centage of  frequency,  347 

—  "  surra  "   in,  95 

—  trypanosomes  in  blood  of,  diseases  asso- 

ciated with,  67,'  68  _ 

Horwood,   polypoid  tumour  of  cervix   uteri 

with  Schistosoma  infection,  643 
Host,  influence  of  parasites  on,  8 
House-fly,   diseases  spread  by,   586 

—  see  also  Musca  domestica 
Howard,  larvae  of  Piophila  casei,  583 

—  Melanolestes  morio,  540 

—  and  Clark,  bug  carrier  of  virus  of  polio- 

myelitis, 536 
Howardia,  characters,  567,  568 
Howardina,  characters,  564 
Huber,  chemically  toxic  effects  of  Ascaridm, 

650 

—  limitations  in  male  fern  administration, 

671 

—  sites  of  cysticercus  in  body,  664 
Hulecoetomyia,  characters,  564 
Human   parasites,  medical   works  on,   617 
Humble-bee,  nests  of,  larvae  of  Homalomyia 

canicularis  found  in,  584 

Hungary,  Simulium  columhaschensis  plague 
in,  578 

Htinsche,  infection  with  Demodex  follicu- 
lorum,  708 

Hyalomma  cegyptium,  characters  and  mor- 
phology,  501,   502 

Crithidia  parasitic   in,   104 


Hyalomma  cegyptium,   farm  stock   sufferers 
from,   in   South   Africa,   502 

—  characters  of,  497 
Hydatid,  see  Echinococcus 

—  disease,  eosinophilia  in,  652 

—  intoxication,  353 

—  sand,  350 

Hydrocele  in  filariasis,  401 
Hydrophobia,   cell  inclusions  in,  207,   208 
Hydrot(Ba  meteorica,  characters,  611 

larvae  of,  habitat,  585 

Hymenolepididce,   309,   323 
Hymenolepis  diminuta,  324,  662 

hosts  of,  326 

intermediate,  327,  328 

—  —  larva  of,  morphology,  328 

—  —  morphology,  326 

occurrence  in  man,  cases  reported,  326 

synonyms,  326 

—  lanceolata,  662 

—  —  hosts  of,  329 

larva,  host  of,  329 

morphology,  328 

synonyms,  328 

—  morphology,   323 

—  murina,   larval   stage,   development   into 

tapeworm,  305 

—  nana,  development,  324 

— ' —  diagnosis  of  presence  in  body,  661 

expulsion  of,  drugs  used  for,  661,  662 

geographical  distribution,  324 

habitat  in  body,   661 

infection    with,    symptoms    following, 

324 

—  —  morphology,  323 

occurrence  in  man,  326 

prevalence  in  children,  661 

c^uestion  of  identity  with  H.  murina, 

324,  325 

—  --  symptoms  following  infection  by,  661 

—  --  synonyms,  323 

—  (Tcenia)    murina,    development    without 

intermediate  host,  17 
larval  stages  occurring  in  rat  flea, 

17 
omission  of  intermediate  host  by, 

326 
Hymenoptera,  characters,  531,  532 
Hypochondriac  region,  right,  liver-fluke  in, 

244 
Hypoderma  hovis  (cattle  or  warble  fly),  595 
invading  human  integument, 

595,  596 

larvae  of,  in  nose,  724 

invading     upper     eyeliH, 

596 
— migration     in     body     of 

cattle,  595 

—  diana,  596 

—  lineata,  596 

geographical  distribution,  596 

Hypotricha,  29,  200 
Hystrichopsylla,  548 

—  distinctive  characters,  545 


864 


THE    ANIMAL    PARASITES    OF    MAN 


Ichneumon,    Indian,    host    of    Paragonimus 

compactus,  251 
Ichthyol  in  chyluria  from  Filaria  bancrofti 

infection,  677 

—  paste,  application  in  creeping  disease,  732 
Ichthyophthirius  multifiliis,  206 

morpholog-y    and    life-history,    207 

Ichthyosporidium,  195 

Ijima,  experimental  infection  of  man  with 
Dibothriocephalus  latus,  312 

—  on   Amoeba   miurai,   46 

—  Tristrongylus  instabilis  in  man,  435 
India,  nasal  myiasis  in,  588 

—  North-West     Provinces,     percentage     of 

pariah  dogs  in,  affected  with  Paropis- 
thorchis   caninus,   257 

Indian  Plague  Committee,  proof  of  infec- 
tion with  Xenopsylla  cheopis,  547 

Infusoria,    198 

—  characters  and  habitat,  29,   198,   199 

—  classification,   199 

—  digestive  processes,  26 

—  encystment  of,    199 

—  hosts  of,  199 

—  macronucleus  and  micronucleus  of,  198 

—  mode  of  life,  199 

—  morphology  of,  198 

—  reproduction  of,  198 

Ingram,  Ehinosporidium  cysts,  197 
Inguinal  tumour,  ascarides  escaping  from, 

656 
Inouye,  lung  fluke  disease,  639 
Insecta,  abdomen  of,  529 

—  anatomy,  529 

—  blood  of,  colourless,  530 

—  development  of,  530 

—  epidermis  of,  530 

—  faceted  eyes  of,  530 

—  head  of,  529 

—  intestinal  canal   of,   530 

—  metamorphosis  of,  530 

—  nervous   system  of,   central,   530 
intestinal,  530 

—  orders  of,  531 

—  organs  of  respiration,  530 

of  touch,    smell   and   hearing,   530 

—  pharyngeal   ganglia  of,   530 

—  sexual  organs  of,  530 

—  sexually  distinct,   530 

—  thorax  of,  529 

Insect   flagellates,    102,    104,    737 
experimentally  introduced  into  verte- 
brates,  104,   112,  737,   738,   739 
Intestinal  canal  of  Insecta,  530 
of  nematodes,  363 

—  obstruction,  Ascaris  in  appendix  causing, 

654 
set  up  by  ascarides,  657 

—  tract,    habitat    of    Oxyuris    vermicularis, 

467 
Intestine,   blood-vessels   of,    penetration   by 
amoebae,  36 

—  coccidiosis  of,  in  man,  cases,  148 

• —  Dipylidium  caninuvi  parasitic  in,  660 


Intestine,    human.    Entamoeba    coli   parasite 
of,  618 

invasion  by  Metastrongylus  apri,  433 

larvae  of  Homalomyia  found  in,  584 

Myriapoda  parasitic  in,  483 

—  large,    cystic    stage    of    Lamblia    intes- 

tinalis  found  in,  59 

Entamoeba  histolytica  present  in,  38 

high  injections  into,  in  evacuation  of 

Oxyuridoe,  698 
human,    Trichuris   trichiura   parasitic 

in,  678 
irrigation    in    gangrenous    dysentery, 

619 

—  larvae  of  Gastrophilus  inhabiting,  599 

—  migration  of  oncospheres  from,  to  liver, 

302 

of  Oxyuris  vermicularis  from,  lesions 

and  irritative  symptoms  set  up  by,  694, 
695 

—  number  of  females  of  Ancylostoma  duo- 

denale   present   in,   mode    of   reckoning 
(footnote),  454 

—  occlusion   of,   due  to  massive   accumula- 

tion  of  ascarides,   657 

—  parasites  of,  in  relation  to  appendicitis, 

views    of    authors    regarding,    652,    653, 
654,  655 

invading  vermiform  appendix,  authors 

recording  cases  of,  652 

—  pathological   changes  m,   due  to  ova  of 

Schistosoma  japonicum,  281 

—  perforation  by  Ascaris,  655,  656 
following   diseased   processes,   656 

—  small,  Ascaris  lumbricoides  parasitic  in, 

687 
Dibothriocephalus   latus   parasitic   in, 

658 
human,    inhabited   by    Tcenia   solium, 

662 

Taenia  saginata  parasitic  in,  667 

inhabited  by   Hymenolepis   nana,   661 

migration     of     Ascaris     lumbricoides 

from,   to  other  parts  of  body,  464 
normal    habitat    of    Ascaris    lumbri- 
coides, 464 
situation     of     Trichomonas     intes- 

tinalis,  55 
possible      invasion     by     Balantidium 

minutum,  204 

—  stenosis  of,  following  infection  by  Tcsnia 

solium,  662 

—  Strongyloides   stercoralis   in,   755 

—  ulceration    of,    associated    with    Balanti- 

dium coli,  202 

—  Yorticellse  in,  206 

—  see  also  Myiasis,  intestinal 

Intra  vitam  staining  of  fresh  preparations 

of  Protozoa,  746 
Inundation  disease,  see  Kedani 
Iodide,  tincture  of,  applications  in  creeping 

disease,  731 
Iodine  enemata  in  flagellate  dysentery,  625 

—  tincture    of,    in    treatment    of    cutaneous 

and  muscular   cysticerci,   663 


INDEX 


865 


Iodoform  with  bicarbonate  of  soda,  adminis- 
tration of,  in  expulsion  of  ascarides,  694 
Ipecacuanha  in  amoebic  dysentery,  619 

—  de-emetinized,  in  balantidian  dysentery, 

637 
Iridocyclitis   in   trypanosomiasis,   623 
Iron-hsematoxylin  stain,  Heidenhain's,  752 
Isaac  and  van  Velden,  dissolution  of  para- 
sitic products  in  serum  of  patients  with 
bothriocephalus  antemia,  645 
Isospora,    149 

—  bigemina,  hosts  of,  149 

morphology,  149 

occurrence  in  man,  149 

synonyms,    149 

Israelites,  ''fiery  serpents"  molesting, 
probable  nature  of,  386 

Itch   mites,   see   Sarcoptidoe 

Itching  resulting  from  clothes  louse  infec- 
tion,  711 

—  set  up  by  Tetranychus  molestissimus ,  488 
Ivers,  infection  with  Demodex  folUculorum, 

708 
Ixodce,  characters  of,  496,  497 
Ixodes,  characters  of,  497 

—  hexagonus,   characters   and   morphology, 

500 

hosts  of,  500 

synonyms,  500 

—  holocychis,  characters,  499 

symptoms  resulting  from  attacks  of, 

499 

—  plumbeus  (dog  tick),  length  of  life  apart 

from  host,  495 

—  reduvius,  act  of  coitus  in,  495 
disinfection   against,   704 

—  —  infection  by,  symptoms,  704 
larvae  of,  495 

life-history  of,  494 

method  of  oviposition,  494 

see  also  Ixodes  ricinus 

—  ricinus  (dog  tick),  bite  of,  effects,  498 
—  prophylaxis  against,  498 

characters  and  morphology,  497,  498 

confusion  in  nomenclature,  499 

geographical  distribution,  499 

habitat  and  hosts  of,  498,  499 

synonyms,  499 

transmission  of  Babesia  bovis  by,  177 

Ixodidoe  (ticks),  carriers  of  various  diseases 
to  animals  and  man,  493 

characters  of,  493 

classification  of,  496 

genera  of,   synonyms,  497 

synopsis  of,  497 

Ixodinoe,  characters  of,  496,  497 

—  transmission  of  Hmmosporidia  by,  704 

—  and   Argantinoe,    distinguishing   features 

between,  505 


Jackal,    Dipylidium   caninum   parasitic    in, 

322 
Jaksch,     von,     causation     of     ancylostoma 

anaemia,  647 


Jamaica,   Amblyomma   cayennense   pest   in, 
501 

—  Margaropus   annulatus  australis   pest  to 

man  in,  505 
James,    genera   of    Anopheliues    (footnotes), 

562,  563 
Jaiiowski  on  the  Trichomonads,  55 

—  presence  of  Cercomonads  in  intestine,  62 
Janthinosoma,   characters,   563,   571 
Japan,    Central,    percentage    of    population 

infected  with  Clonorchis  endemicus ,  260 

—  illness  set  up  by  kedani  mite  in,  487 
Japanese    river   or   inundation    disease,    see 

Kedani 
Jaundice,  malignant,  in  dogs,  carrier  of,  493 

cause  of,   177 

Jaw,  upper,  nematode  larvae  in  periosteum 

of,  associated  with  gingivitis,  378 
Jejunum,    flagellate    stage    of    Lamblia    in- 

testinalis   found  in,  59 

—  human,     habitat    of    Ancylostoma     duo- 

denale,  450 
Jerboa,   haemogregarine   in,   154 

—  inoculation  of  Trypanosoma  lewisi  into, 

90 
Jews,  infection  with  Taenia  saginata,  340 

—  inoculation  against  Oriental  sore,  108 
Jigger,  see  Dermatophilus  penetrans 
Joblotina,  characters,  565 
Johannseniella,  579,  580 

Johns,  cultivation  of  malarial  parasites,  170 
Jungklauss's  preparation  as  vermifuge,  673 
Jiirgens,  case  of  amoebae  in  urine,  46 

—  intestinal    amoebae,    36,    37 


Kabyles,  tamne  or  thimni  of,  598 
Kahane,    earth-eating    in    connection    with 
Trichuris  trichiiira  infection,  679 

—  Trichocephali  in  appendix,  655 

—  trichocephalus  anaemia,  651 
Kala-azar,  canine,  similarity  to  human   in- 
fantile form,  103 

transmission  by  dog  fleas,  103 

—  dissemination  of  bug  possibly  connected 

with,   107,  536 

—  Indian,  agents  of  transmission,  107 

geographical  distribution,  105 

incubation  period,  626 

mortality   great,   626 

parasite  of,   105,   626 

preventive  measures,  626,  627 

symptoms  and  course,  626 

treatment,  626 

—  infantile,    109,   627 
aetiology.  111,  627 

geographical  distribution,  109 

in  adolescents  and  adults,  109 

natural   infection   of   dogs   with,    110 

preventive  measures,  627 

relation  to  Indian,  109 

—  to    Oriental     sore     shown     experi- 
mentally,   109 
symptoms,  627 


866 


THE    ANIMAL    PARASITES    OF    MAN 


Kala-azar,    infantile,   transmission   by   fleas, 

experiments    to    prove.    111 
treatment,  027 

—  possible  transmission  by  bed  bug,  107,  713 
Kaldrovils,     cysticercus     of     eye     mistaken 

for  foreign  body,  664 
Kamala   as  vermifuge,   673 

—  in  evacuation  of  Oxyuridoe,  697 
Kaposi's  uaplithol  ointment,  application  in 

scabies,   707 
Kartulis,  case  of  amoebae  in  urine,  46 

—  cerebral    abscesses   in    amebiasis,   35 

—  discovery  of  amoebae  in  stools  of  dysen- 

tery  patients,   30 

—  Entamoeba   kartulisi,  44 

—  experiments       proving       connection       of 

amoebae  with  dysentery,  30 

—  Sarcosporidia  in  man,   194 
Karyolysus,  154 

Kautsky,  billiarziasis,  641 

Kayser,  eye  affections  due  to  Liicilia  macel- 

laria,   721 
Kedani      (Japanese      river      or     inundation 

disease),  487,  703 

— prophylaxis  against,  703 

^  _  _  symptoms,  703 

—  mite.  487 

characters  of,  487 

Kelly,  bilharziasis  of  appendix,  642 
Kent,  Herpetomonas  and  Leptomonas,  102 
Kerosene,  destruction  of  Tabanidoe  by,  601 
Kerteszia,   characters,   562,   569 
Kholodkowsky,     post-mortem     discovery     of 

Opisthorchis  felineus,   253 
Kidney  and  liver  cells,  yellow  pigment  in, 

in   ancylostomiasis,   647 
Kinghorn,     transmission     of     Trypanosoma 

rhodesiense,  69 

—  and   Yorke,   tsetse-fly   transmitting    Try- 

panosoma rhodesiense,  608 

transmission  of  Trypanosoma  rhode- 
siense, 81 

Kirmisson,  trichocephalus  infection  in  re- 
lation to  appendicitis,  653 

Klebs,  early  researches  on   malaria,   156 

Klencke,  early  mention  and  depiction  of 
malarial  parasites,   157 

Kloss,  researches  on  Coccidia,  136 

Klossia,    141 

Knackers'  yards,  infection  of  rats  with  Tri- 
chinella   in,   427 

Knoch,  J.,  views  on  development  of 
cestodes,  16 

Kobayashi,  second  intermediate  host  for 
Clonorchis  endemicus ,  261 

Koch,  R.,  artificial  infection  of  species  of 
Glossina  with  human  trypanosome,  605 

discovery     of     amoebae     in     stools     of 

dysentery   patients,  30 

investigations      of     Proteosoma      and 

Halteridium   in   birds,    158 

relapses      and      latent      infection      of 

malaria,  158 

researches  on  malaria,  158 

on  SpirochcBta  duttoni,  117 


Koch's  blue  bodies  in  Theileria  parva,  179 
Kolliker,  investigation  of  gregarines,  129 
Koneff,    favourable    effects    of   expulsion    of 

Ascaridce,  650 
Kousso  flowers  as  vermifuge,  673 

—  in  evacuation  of  Oxyiiridw,  697 
Kraft,  filmaron  as  vermifuge,  672 

Kruse      and      Pasquale,      nomenclature      of 

amoebae,    31 
Kiichenmeister,  F.,  experimental  rearing  of 

tapeworms,   15 
experiments    as    to    metamorphosis   of 

tapeworms,  15 

expulsion  of  Ascaridm,  693 

nature   of  cysticerci,  282 

Kuhnt,  infection  of  human  eye  with  fllaria, 

406 
Kummerfeld's  wash  for  clothes  lice,  616 
Kunstler,  genus  Giardia,  736 
Kurlow,       blood-stained       diarrhoea       from 

Strongyloides  stercoralis  infection,  675 
Kutner,    favourable    effect   of   expulsion    of 

Ascaridce,  649 

—  treatment  of  bilharziasis,  643 


Labadie-Lagkave    and    Deguy,    invasion    of 
lymphatic    vessels    by    Onchocerca    vol>- 
vulus,  418 
Labbe,  copulation  in  Coccidia,  137 
Lacompte,  nematodes  in  human  eye,  412 
LoRlaps    echidniniis,   Leishmau    granules    in, 
493 

—  stabularis,  493 

Lafleur,  see  Councilman  and  Laflenr 
Lagocheilascaris,  characters,  466 

—  minor,   host  of,  467 
lesions  set  up  by,  467 

—  —  morphology,  467 

Lakes,  mosquitoes  depositing  ova  in,  553 
Lama,  possible  carrier  of  leprosy,  613 
Lambkin's  mercury  cream  in  treatment  of 

syphilis  prevailing  in  Uganda,  632 
Lambl,      discovery     of     human      intestinal 

amoebae,   29 
Lamblia    intestinalis ,    57,    625,    736 

association  with  diarrhoea,  59,  60,  625 

treatment,    625 

—  —  characters,   57 
flagella   of,   57,   58 

—  —  hosts    of,    59 

—  —  infection  with,  60 
nuclear   apparatus,   58 

site    in    intestine    of    flagellate    and 

cystic   stages,    59 

synonyms,  57,  736 

Lankester,  liver-fluke  in  abscess  of  ear,  244 

—  Sarcocystis,    193 
Lankesterella,  154 
Larva  migrans,  599 

Larvae,  dipterous,  in  conjunctiva,  716 

—  —  in   nasal   accessory   sinuses,   717 
in  nose  in  enormous  numbers,  716,  717 

—  in   wounds,   movement  of,   723 


INDEX 


867 


Larvae  in  wounds,  see  also  under  Names  of 
Parasites 

Larvicides,  use  in  campaign  against  mos- 
quitoes, 636 

Larynx,  ascarides  invading,  691 

—  leeches   in,   699,   700 

La  Spada,  echinococcus  of  liver  rupturing 
into  abdominal  cavity,  652 

Lasioconops,  characters,  564 

Lassar's  paste,  application  in  creeping 
disease,  732 

Laurer's  canal  of  trematodes,   221,  222 

Laveran,  A.,  classification  of  trypanosomes, 
71 

cross-immunity  experiments  with  Try- 
panosoma rhodesiense  and  T.  hrucei,  80, 
94 

—  with   trypanosomes,   80 

discovery    of  true   malarial   parasites 

by,   157 

—  —  latent  forms  of  trypanosomes,  74 

on  Leucocytozoa,   153,  742 

Trypanosoma  pecaudi,  95 

and  Franchini,  inoculation  experi- 
ments with  Crithidia  fasciculata,  104 

with     Herpetomonas     cteno- 

cephali,   103 

—  — with    H.    pattoni,    103 

and  Mesnil,   isolation   of  sarcocystin, 

191 

—  —  —  on       the       spore       of      Sarcocystis 

tenella,    193 

"  Trypanosomes  et  Trypano- 
somiases,"     617 

and   Thiroux,    treatment    of    sleeping 

sickness,  623 

Laverania,  characters,   164,   569 

—  nialarice        (Plasmodium        falciparum), 

crescents  of,   162,   167,   168 

sites   of   development,    169 

—  —  cultivation  of,  clumping  in,  172 

—  cultures    of,    number    of    spores 

produced,  172 

development,  duration  of,  167 

distinctive  characters  of,  169 

—  —  invasion  of  spleen  by,  168 

—  —  —  —  merozoites,   number  of,   168 
number   in   one   red   blood   cor- 
puscle, 167 

oocysts  of,  in  stomach  of  Ano- 
pheles, 163 

ookinete     of,     in     stomach     of 

Anopheles  maculipennis,   162 

parasite    of    malignant    tertian 

or  sub-tertian  fever,  167 

and      quotidian 

malaria,    167,    633 

—  ^ pathological  effects,  634 

question  of  varieties  or  sub- 
species, 167 

"  signet-ring  "  stage,  167 

sporozoites,   169 

stages  of  development  in  intes- 
tine of  Anopheles  maculipennis,  162 

—  —  synonyms,   167 

trophozoites  of,  167,  168 


Leo,  R.  J.,  creeping  disease,  729 
Leeches  in  upper  air  passages,  699,  700 

cases       reported       by       various 

authors,  699,   700,  701 

mention  among  ancient  writers, 

699,  700 

—  invading  body,   means  of  riddance,  701 

—  see   also   Hirudinea,   Rhyncobdellidos 
Leeuwenhoek,  opposition  to  theory  of  spon- 
taneous generation,   10 

Leger,  L.,  classification  of  Coccidiidea,  141, 
142 

—  genus  Crithidia,   104 

—  researches  on  Coccidia,  137 

Leger,  M.,  proportion  of  population  in 
Tonkin  infected  with  Glonorchis  en- 
demicus,  260 

Leger,  M.  and  A.,  proposed  classification  of 
Leucocytozoa,  153 

Leichtenstern,  bothriocephalus  anaemia,  646 

—  toxic    symptoms    following    thymol    ad- 

ministration, 686 
Leidy,  genus  Endammha,  31,  34,  734 
Leignathus  sylviarum,  493 
Leiper,  R.  T.,  Gastrodiscoides,  236 
host  of  Filaria  loa,  601 

—  —  identity  of  (Esophagostomum  hrumpti 

with   (Es.   apiostomum,   444 

report  of  Bilharzia  Mission  under,  277 

Leipzig,    frequency   of   infection   of   various 

organs    of    animals    with    echinococcus 

slaughtered  at,  347 
Leiseri  ng,  percentage  of  rats  infected  with 

Trichinella,  427 
Leishman,     Sir    W.     B.,     experimental    re- 
searches   on    infection    with   Spirochceta 

duttoni,   117,    118 
on    parasite    of    Indian    kala-azar, 

105 

treatment  of  Indian  kala-azar,  626 

Leishman-Donovan    body,    see    Leishmania 

donovani 
Leishman    granules    in    Laelaps    echidninus, 

493 
Leishmania,  67,  104  ' 

—  donovani,  105 

cause   of  Indian   kala-azar,   105,   626 

cultivation  methods,  106 

inoculation  experiments  with,  107 

localization  of  infection,   105 

morphology,   105,  106 

possible  mode  of  transmission,   107 

—  evolution    from    flagellates    of    inverte- 

brates, 739 

—  infantum.,   cause   of   infantile   kala-azar, 

105,  109,  627 

cultivation  methods,   109 

immunity  to,  112 

in  dogs,  110 

—  —  inoculation,    110 

animals   suitable   for,   110 

probable    transmitter.    111 

—  probable   origin   of,   103,   739 

—  tropica,  105,   107 

cause  of  Oriental  sore,  105,  107,  627 

i cultivation  methods,   108 


868 


THE   ANIMAL    PARASITES    OF    MAN 


Leishmania  tropica,  hosts  of,   108 

inoculation,  experimental,  108 

possible  transmitters,    108 

synonyms,   107 

Leishmaniasis,  cutaneous,   107 

—  dermo-mucosal,  supposed  mode  of  trans- 

mission   in    Paraguay,   739 

—  evolution  of,  relation  of  experimental  in- 

troduction of  insect  flagellates  into  ver- 
tebrates on,   737,   738,   739 

—  experimental   production   in    white  mice, 

103 

—  geographical    distribution,    105,    107,    109 

—  infantile,  see  Kala-azar,  infantile 

—  naso-oral,  see  Espundia 

—  possible  reservoirs,  738,   739 

—  treatment,    G26-629 
Leishman's  stain,  750 

Lemaire,    herpetomonad    flagellate    in    cul- 
tures of  blood  and  organs  of  gecko,  739 
Lenhartz,  bothriocephalus  anaemia,  646 
Lentospora  cerebralis,  184 
Lepidoptera,  characters,  531,  532 
Leprosy,  possible  carrier  of,  579,  613 
Leptidce,   603 

—  blood-sucking  species,  603 

—  characters,   603 
Leptis  scolopacea,  603 

—  strigosa,   603 
Leptodera,  life-history  of,   19 

—  appendiculata,  occasional  parasite,  7 

—  pellio,  facultative  parasitism  of,  8 
Leptomonas,   102 

—  biitschlii,   102 
Leptotheca,  184 

Leptiis  autumnalis  (grass,  harvest  or  goose- 
berry mite),  animals  attacked  by,  486 

skin  irritation  set  up  by,  702 

habitat  of,  485 

hosts  of,  485,  486 

—  ■ —  nut  and  fruit  pickers  affected  by,  485 

skin   affection  set  up  by,  485,  486 

so-called  proboscis  of,  485,   486 

—  geographical  distribution  of  species,  486 

—  undescribed  species  of,  486 

Lesbini,    dipterous    larvae    in    nose    in   enor- 
mous numbers,   717 
Letulle,  pathological  changes  in  rectum  due 

to  Schistosoma  hosmatohium,   274 
Leuckart,    E..,    advances    in    helminthology 

due  to,  15,  16 
attempt  at  self-infection  with  Ascaris 

lumhricoides,  464,  465 

change  of  host  in  parasites,  20,  21 

classes  of  parasites,   1 

development    of    Acanthocephala    and 

TAnguatulida,  17 

, of  alveolar  echinococcus,  357,  358 

of  nematodes,    17 

—of  Trichinella  spiralis,  423 

distinction    between    Cercomonas    and 

Trichomonas,   54 
experimental         self-infection         with 

Oxyuris    vermiciilaris ,   469 
facultative  parasitism,  7 


Leuckart,     K.,     feeding     experiments     with 

T(Bnia  saginata,  340 
with  Trichinellae,  423 

—  —  growth  of  echinococcus,  354 

—  —  heterogony     in      Strongyloides      stei^- 

coralis,  381 

—  —  method    of    infection    with    Trichuris 

ovis,  420 

migration  of  oncospheres,  302 

name  of  Coccidia  first  given  by,  135 

Trichocephalus     in     association     with 

cholera,  658 
and  Thomas,  P.,  life-history  of  liver-. 

fluke,   241 
Leucocytogregarina,   154 

—  canis,   life-cycle   diagram,    155 

transmission  from  dog  to  dog  by  tick, 

155 
Leucocytogregarines,  154 
Leucocytosis  in  bilharziasis,  642 
Leucocytozoa,  action  of,  on  red  blood  cells, 

153,   742 

—  classification   proposed,   153 

—  hosts  of,  153 

—  morphology    of,    153 

—  schizogony  of,   153,  742 
Leucocytozoon  type  of  Iloimosporidia,  152 

—  lovati,  schizogony  in,  153 

—  ziemanni,  schizogony  in,  153 
Leucomaines,  effects  on  living  organisms,  9 
Levaditi,   cultivation   of   spinal   ganglia   of 

rabid  monkeys,  210 
Lewandowsky,  infection  with  Demodex  fol- 

liculorum,   708 
Lewin,  expulsion  of  Ascarides,  693 
Lewis,  finding  of  intestinal  amoebae,  29 

—  studies  of  filariasis,  391 
Leydenia,  49 

—  gemmipara,   49 
in  ascites,  49 

association  with  possible   ascites  and 

malignant  growth  in   abdomen,  49,   50 
characters  of,  49 

—  —  cytoplasm  containing  blood  corpuscles, 

50 
pseudopodia    of,    joining    several    in- 
dividuals, 49 
Leydig,  psorosperms,   181 
Lice,  Ilerpetomonads  in  gut  of,  103 

—  transmission   of   relapsing   fever   by,    120 

—  wingless,  owing  to  parasitic  life,  3 

—  see  also  Pedicididce 

Lieberkiihn,  investigations  of  Coccidia,   135 
of   gregarines,    130 

—  psorosperms,   181 

Liesen,  Ascaris  in  peritoneal  cavity,  656^ 
Ligula,      excretory      apparatus,      collecting 
tubes,  island  formation,  292 

—  plerocercoid  of,  300 
Limatus,   characters,  565 

Limnaeus,  species  other  than  L.  triincatulus 
intermediate  hosts  of  Fasciola  hepatica, 
242 

—  truncatulus,    amount    of    ova    deposited 

by,  242 
geographical  distribution,  241 


INDEX 


869 


Limnoeus  truncatidus,  intermediate  host  of 
Fasciola   hepatica   (?),  240,   241 

hosts  of  liver-fluke,  240,  241 

Limnatis,    characters,    482 

—  nilotica,  characters,  482 

geographical  distribution,  482 

habitat,  482 

only   leech   of   clinical    importance    as 

parasite,   699,   701 

synonyms,  482 

Lindblad,  Dipylidium  caninum,  660 
Lindner,  G.,  peritrichal  Infusoria  (stalkless 

Vorticella),   206 
Lindsay,     possible    mode    of  -transmission 

of  dermo-mucosal  leishmaniasis  to  man 

in   Paraguay,   739 
Linguatula,  523,  524 

—  rhinaria,  characters  and  morphology,  524 
development   and   life-history  of,   524, 

525,  526 

larvae  of,  524,  525 

occurrence   at   autopsies,   526 

organs  of  body  invaded  by,  524,  525, 

526 

ova  of,  524,  525 

parasitic  in   nasal   cavity  of  animals 

and   man,   523,   524 
synonyms,    524 

—  serrata,  hosts  of,  527 

synonyms   (footnote),  527 

Linguatulida,   development  of,   17 
Linguatulidoe,  blood-sucking,  523 

—  change  of  original  features  in,  4 

—  characters  and  morphology,  523 

—  hosts  of,  523 

—  larvae  of,  523 

—  nature  of,  2 

—  relation  to  Arachnoidea,   19 

—  separation  from  Helminthes,  2 

Lini,  escape  of  Ascarides  from  umbilicus, 
656 

Linnaeus,  discoveries  as  to  origin  of  Hel- 
minthes,  10,   11 

—  so-called     dysentery     infection     due     to 

mites,  512 
Lipari,   cysticerci   of  brain,   664 
Lipuria  in  bilharziasis,  641 
Lithocystis,  endoplasm  of,  contents,   131 
Liver,  abscess  of,  association  of  Entamoeba 

histolytica  with,  35 

of  Noc's  entamoeba  with,  41 

caused  by  invasion  of  Ascaridoe,  690 

due  to  amoebic  dysentery,  treatment, 

620 
set  up  by  amoebae,  35 

—  and     bile-ducts,     habitat     of     Clonorchis 

endemicus,   259,   260 

—  and  kidney  cells,  yellow  pigment  in,  in 

ancylostomiasis,  647 

—  and    portal    vein.    Schistosoma    hcRmato- 

hium,  most  easily  found  post  mortem  in, 
273 

—  coccidiosis  of,  in  man,  cases,  148 

—  encystment    of    Porocephalus    constrictus 

in,  526,  527 

—  female  Ascarides  depositing  ova  in,  689 


Liver,  human,  eggs  of  Schistosoma  japoni- 
cum,  showing  "  spines  "  and  "  hoods  " 
at    opposite   pole,   279 

—  invasion  by  larvae  of  Linguatula  rhinaria, 

525,  526 

—  migration  of  oncospheres  from  intestine 

to,  302 

—  pathological  changes  associated  with  in- 

vasion by   Opisthorchis  felineus,  253 

—  —  —  in,    due    to    ova    of    Schistosoma 

japo7iicum,  281 
set    up    by    Clonorchis    endemicus , 

260 
Liver-fluke,  supposed  origin  of,   10 

—  see  also  Fasciola  hepatica 

—  disease,  diagnosis,  242 

in  man,  242 

in  sheep,  238 

ravages  caused  by,  238,  239 

stages  of,  240,  241 

pathological  anatomy,  241 

symptoms,  239 

Liverpool  School  of  Tropical  Medicine,  ex- 
pedition to  investigate  trypanosome  in- 
fections,  68 

Lizards,  haemogregarines  from,  154 

Loa,  morphology,  409,  411 

— •  loa,  duration  of  life  of,  414 

early  historical  accounts  of,  412 

geographical  distribution,  414 

larvae  of,  in  blood,  412,  414 

periodicity,  413 

structure,  412 

lesions  produced  through  invasion  by, 

413,  414 

life-history,  414 

morphology,  409,  411 

ova  of,  410 

•  sites  of  body  invaded  by,  412,  678 

synonyms,  409 

Lobaczewski,  prophylaxis  against  body, 
head   and  clothes  lice,  615 

Lobker,  cause  of  ancylostome  anaemia,  648 

Locusts  injurious  to  man,  542 

Looss,  infection  by  Ancylostoma  duodenale 
through  skin,  683 

—  origin  of  lateral-spined  eggs  of  Schisto- 

soma  hcematohium,,  273 

—  prevalence    of    Heterophyes    heterophyes, 

264 

—  skin    affections    set    up    by    invasion    of 

larvae  of  Anciflostoma  duodenale,  455 

—  symptoms  of  lymphangitis  from  Filaria 

bancrofti  infection,  676 

—  toxic  action  of  ancylostomes,  647 

—  Trichostrongylus  instabilis  in  man,  435 
Lophius  piscatorius,   186 
Lophoscelomyia,   characters,   562,   568 
Losch,    discovery    of    intestinal    amoebae    in 

case    of   dysentery,    29,   30,   32 
Loschia,  34 

Lota  vulgaris,  see  Burbot 
Lounsbury,       life-cycle       of       Amblyomma 

hebrceum,  495 
Louse  disease,  historical  instances  of  death 

from,  711 


870 


THE   ANIMAL    PARASITES    OF    MAN 


Low,   personal   experiments   with   regard  to 
malaria  infection^  158 

—  treatment  of  Oriental  sore,  628 
Lucilia   argyrocephala   cause   of   myiasis   in 

French  West  Africa,  G14 
— -  ccesar,  588 

—  macellaria,      larvae      of,       causing      eye 

diseases,  721 

■  —  in  nose,  715,  716 

see    also   Myiasis,   nasal 

on  cutaneous  surface,  721,  722 

penetrating  auditory  meatus,  721 

—  nobilis,    larvae    (maggots)    of,    discharge 

from  auditory  meatus,  588 

—  sericata,  588 
Lumbricosis,  typhoid,  650 

Lumbricus,  Monocystis  agilis  from  seminal 
vessels  of,   130,   132 

—  teres,   see   Ascaris    lumhricoides,   464 
Lund's  larva,  characters,  593 

Lung,  abscess  of,  set  up  by  amoebae,  35 

—  amoebae   found   in,   45 

—  Balantidium   coli  occurring  in,   202 

—  bilharziasis  of,  642,  643 

—  gangrene     of,     possible     occurrence     of 

Cercomonads  in,  62 

—  invasion  by  Fasciola  gigantica,  245 

by   Paragoninius    ringeri,   251 

by  Schistosoma  hmmatohium,  274 

—  Trichomonads  found  in,  56 
Lung-fluke    disease,    geographical    distribu- 
tion, 639 

prognosis,  640 

symptoms,  639 

treatment,  640 

Lussana,      toxic      theory      of      ancylostome 

anaemia,  646 
Liitz,  ascarides  in  pulmonary  artery,  656 

—  Ceratopogoninoe  described  by,  580 

—  experimental  infection  with  Ascaris  hrm- 

hricoides,  465 

—  favourable      effects      of      expulsion      of 

Ascaridm,  649 

—  perforative  peritonitis  due  to  Ascaris,  656 
Lyctocoris  campestris,   bite   of,  541 

characters,    541 

Lygceidce,  characters,  541 
Lymphangitis  from  Filaria   bancrofti  infec- 
tion, symptoms,  676 

—  in  filariasis,  401 

Lymphatic    glands,    enlarged,    in    filariasis, 
402 

—  vessels   and   glands,   destruction   without 

lymphatic    obstruction,    401 

distribution    and    connections    of, 

400,  401 

—  vessels,  invasion  by  Onchocerca  volvulus, 

418,  419 
Lymphatics,  Strongyloides  stercoralis  in,  755 
Lynch,  human  trichomoniasis,  734 
Lynchia,  transmitting  Haltcridium ,  151 
Lyperosia,    differentiation    from    Stomoxys, 

610 

—  exiqua,  life-history,  610 

—  irrifans,  var.  weisii,  610 


Macaciis    sinicus,    inoculation    with    Leish- 

mania  donovani,   107 
MacCallum,   "  exflagellation,"   152 

—  investigations    of    Proteosoma    and    Hal- 

tcridium   in    birds,    158 

McDonagh,  J.  E.  R.,  life-cycle  of  organism 
of  syphilis,   124 

MacFadyean  and  Stockman,  Babesia  diver- 
gens,  177 

Macfie,   Trypanosoma  nigeriense,  76 

—  and    Gallagher,    treatment    of    sleeping 

sickness,  622 
Mackenzie,   periodicity  of  larvae  of  Filaria 

bancrofti  in  peripheral  blood,  393 
Mackie,  suggested  transmission  of  relapsing 

fever   by   lice,    120 

—  treatment  of  Indian  kala-azar,  623 
Macleayia,   characters,  563 
MacNeal,  see  Novy  and  MacNeal 
Macrostoma  niesnili,   57,   735 

Maculae  caerulae  (taches  bleties)  due  to  in- 
fection  by  crab   louse,   712 

Maggots,  see  under  Names  of  Parasites  and 
Regions  of  Body 

—  in  nose,  see  Myiasis,  nasal 
Magnesium    sulphate    in    flagellate    dysen- 
tery, 625 

Maillard,  fatal  cases  of  nasal  myiasis,  718 
Majochi,     case     of    intertrigo     set     up    by 
Oxyuris  vermicularis ,  696 

—  infection  with  Demodex  folliculorum,  708 
Mai     de    caderas    in    horses,    trypanosomes 

associated    with,    68 

trypanosome   causing,   96 

Malaria,   acute,   156 

—  atypical  forms,   634 

—  campaign     against,    commencement    and 

progress  of,  158 

—  chronic,  156 

—  development  of  parasites  of,   159 

—  diagnosis   (pathognomonic  signs),  635 

—  geographical  distribution,  155 

—  historical,    157 

—  in  birds  spread  by  Culex,   158 
— -  in  man,   155 

—  latent,  in  children  of  natives,  158 

—  masked,  156 

—  parasites  of,  164-170,  633 

asexual     generation,     cultivation     in 

vitro,   170 

copulation,   160,   161,    162 

exflagellation    (footnote),    162 

—  —  gametocytes  of,   160,   161,   162 

human,  development,  159 

occurs  only  in  Anopheles,  158, 159 

differential  characters,  171 

species   of,   164,   633 

—  see     also     Laverania     malarioe, 

Plasmodium  malaria?,  Plasmodium  re- 
lictnm,   Plasmodium    vivax 

macrogametes  of,   160,   161,   162 

—  —  merozoites  of,   159,   160 

—  —  methods  of  detecting,  747 

microgametes  of,   160,    161,   162 

movements,  discovery  of,  157 


INDEX 


871 


Malaria,   parasites   of,   not   transmissible  to 
mammals,   159 

ookinetes,  160,  161,  163 

schizogony  of,  161,  172 

sporozoites  of,    159,    160,   164 

penetration      of     red     blood     cor- 
puscles by,  159,  160 
sporulation,   160,   161,   163 

—  pig-mentation   of   organs,    165    (footnote), 

634 

—  prevention  of  constipation  during,  635 

—  preventive  measures  against  mosquitoes, 

635,  636 

against   parasite   in   man,   635,   636 

by  quinine  administration,  636 

—  prophylaxis,  636,  637 

—  relief  of  symptoms,  635 

—  symptoms,    156,    633 

—  synonyms,   155,  633 

- —  tertian,  malignant,  paroxysms  of,  634        1 

—  treatment  by  quinine,  635 
Malarial  fever,  quartan,  156 

duplex   or   triplex,   appearance   of, 

167 

parasite  of,   166 

malignant  or  sub-tertian,  parasite  of, 

167 
pernicious   symptoms,   explanation, 

172 

quotidian,    156 

rhythmical,  course  of,  155 

symptoms,  155,  633 

tertian,   156 

■  —  simple  or  spring,  parasite  of,  164 

—  —  typical,   clinical  features,  633,  634 
Male  fern,  administration  to  children,  671, 

672 

emulsion,   injection  of,   671 

ethereal    extract    best    vermifuge    for 

TcBTiia  saginata,  670 
dosage  and  method  of  adminis- 
tration, 670,  671 
■ extract  of,  expulsion  of  Hymenolepis 

nana  by,  661 

in  bilharziasis,  643 

in   expulsion   of   ancylostomes,   686 

—  in   intestinal   myiasis,   728 

poisoning,  670,  671 

— antidotes  to  and  remedies  for,  671 

bad  effects  on  vision,  670 

Malignant   malarial    parasites,   sporulation, 

influence  of  temperature  on,  163 

stages  of,  163 

Mallory's   bodies,    208 

Mamma,  tumours  of,  association  of  Diocto- 

phyme  gigas  with,  431 
Mammals,    human    malarial    parasites    not 

transmissible  to,   159 

—  leucocytogregarines   in,    154,    155 

—  r;  d    blood   corpuscles   of,    Babesia   para- 

sitic in,   154 
M:rT,   incidental   parasites   of,  7 

—  infection  with  animal  trypanosome,  96 

—  parasites  found  only  in,  6 
Mange,  see  Dog  mange 


Mangold,   feeding  experiments  with  Taenia 

from  multilocular  echiriococcus,   358 
Manguinhosia,  characters,  562,  568,  569 
Manson,      Sir      Patrick,      development      of 

Paragonimas  ringeri,  251 
discovery    of   Sparganum   7nansoni, 

317 
infection    of    skin    by    Filaria   per- 

stans,    378 

on  Spirochaeta  carteri,  631 

pathognomonic    signs    of    malaria, 

635 
—  prophylaxis       against       ancylosto- 

mia,sis,  685 

researches  on  malaria,  158,  635 

studies  of  filariasis,  391 

treatment  of  Indian  kala-azar,  626 

^  —  of  Oriental  sore,  628 

Manson,   P.    T.,    infected   with   malaria   by 

infected  mosquitoes,  158 
Manson ia,  577 
Manson's     method     of     administration     of 

atoxyl  in  sleeping  sickness,  622 
Manteufel,  immunity  of  Ornithodorus  mou- 

bata  against  infection  with  Spirochceta 

duttoni,   119 
Marchiafava,    discovery    of    movements    in 

malarial   parasites,    157 
Marchoux,    amoebic    abscesses    in    liver    of 

experimental   cats,    35 

—  Spirochceta  gallinarum,  119 

—  and  Couvy,  Leishman  granules  in  Loelaps 

echidninus,  493 
Mareo,  Helminthiasis  meningitiformis ,  649 
Margaropus   annulatus    australis,   hosts   of, 

505 
pest  to  man   in  Jamaica  in  larval 

stage,  505 

—  characters  of,  497 

—  microplus ,   505 

Marx,  male  fern  administration,  671 

—  toxic   action   of  male   fern,  670 
Marzinovsky,    prophylaxis    against    Pedicu- 

lus  vestimenti,  616 
Mastigophora,   28,   50,   760 

—  aggregation  rosettes  of,   51 

—  characters    and   habitat,   28 

Mathis,    carriers    of   Entamoeba   histolytica, 
40 

—  diarrhoea    due    to    Lamblia    intestinalis , 

625 

—  Lamblia  intestinalis ,  59,  60 

—  modification    of    Novy-MacNeal    medium, 

744 
Maurer's  dots,  168,  170,  171 
Maxillary  sinus,  Scolopendra  in,  721 
Mayer's  glychsemalum,  751 

—  hsemalum,   751 
Mbori  in  dromedaries,  06 
Measles,  207 

Meat    inspection,    decrease    of    cysticerci    in 

pork  effected  by,  334 
Meatus,  auditory,  larvae  of  Anthom,yia  plu- 

vialis  found  in,  584 

Rhinosporidium    in,    196 

maggots  of  Lucilia  nobilis  in,  588 


872 


THE   ANIMAL    PARASITES    OF    MAN 


Meatus,    auditory,    synonyms,   438 
Mecistocirrus,  habitat,  438 

—  morphology,  438 

—  fordi,  morphology,  438,  439 

• synonyms,  438 

Medullary  layer  of  Cestoda,  289 
Megarhinince,  characters,  563,  570 
Megarhinus,  characters,  563,  570 
Megnin,  development  of  cestodes,  16 
Mehlis,    discovery    of    progeny    of   Distoma, 

Typhloccelum   flavum-,    and    Cathoemasia 

hians,  12 
Mehlis'  gland  secretion  in  trematodes,  223 
Melanoconion,     distinguishing     characters, 

564,   576 

—  atratus,  characters,  576 

geographical  distribution,  576 

Melanolestes  abdominalis,  540 

—  morio,  geographical  distribution,  540 
synonyms,  540 

Mello-Leitao,    flagellate    dysentery    in    chil- 
dren,  56,  624 
Melnikow-Raswedenkow,      development      of 

alveolar  echinococcus,   357,   358 
Melophagus  ovinus  (sheep  ked),  bite  of,  611 

Crithidia  inhabiting,   104 

Meningitis,   fatal,   peenash   terminating   in, 
716 

—  symptoms  of,  due  to  Ascaridce  infection, 

649 

—  terminating  nasal  myiasis  fatally,  718 
Mense,  expulsion  of  Guinea  worm,  676 
Mercier,  nematodes  in  human  eye,  412 
Mercury,    benzoate    of,    in    infantile    kala- 

azar,  627 

—  cream  (Lambkin's)  in  syphilis  prevailing 

in   Uganda,   632 

—  in  expulsion  of  Strongyloides  stercoralis , 

675 
Mermis,  469 

—  hominis  oris,  469 
Mermithidce,  469 

—  characters,  375 
Merogregarina,   135 
Merogony,   185 
Meront,  185 

Merozoites  of  Coccidiidea,  138,  139,  140 

—  of  malarial  parasites,  161 
Mesenteric  vein,  superior,  tributary  of  por- 
tal vein,  272 

Mesnil,  on  Actinomyxidia,   187 

—  on  Hsemosporidia,  742 

—  on  Haplosporidia,   194 

—  and      Ringenbach,      cross-immunity     ex- 

periments with  trypanosomes,  80 
trypanolytic  reactions,  80 

—  see  also  Laveran  and  Mesnil 
Messineo,   effects   of  experimental   injection 

of  extracts  of  Taenia,   648 
Metagonimus,  264 

—  (Yokogawa   yokogawai,   264,   753 

geographical    distribution,    265 

habitat,    265 

host    and  intermediate  host,  265 

life-history,    265 

morphology,    264 


Metastrongylinae,  characters,  432 
Metastrongylus,  morphology,  432 

—  apri,   hosts  of,  433 

in  man,  cases  recorded,  433 

invasion  of  air-passages  by,  433 

morphology,   432 

synonyms,   432 

Methyl   green,   752 

Methylene  blue  in  bilharziasis,  643 

in   flagellate   diarrhoea,   625 

Metorchiince,  261 

—  morphology,  232 
Metorchis,   261 

—  conjunctus,  organs  of,  diagram  showing, 

258 

—  truncatus ,  habitat  and  hosts  of,  262 
morphology,  261,  262 

organs  of,  diagram  showing,  262 

Metschnikoff,  intestinal  parasites  in  relation 
to  appendicitis,  652,  653 

—  prophylaxis  against  oxyuriasis,  697 
Meyer,  disturbances  of  vision  in  male  fern 

poisoning,  670 
Mibelli,    infection    with    Demodex    follicu- 

lorum,  708 
Mice,  experimental  infection  with  herpeto- 

monads,  103,  104,  112,  737,  738,  739 

with  Sarcocystis  muris,   191 

— with  SpirochoBta  duttoni,  117 

—  natural  herpetomonads  in,  738,  739 

—  occasionally       hosts       of       Hymenolepis 

diminuta,  326 

—  Sarcosporidia  in,  187 

—  spherical    contracted    forms    of    Tricho- 

monas intestinalis  in,  56 
Michelson,    case    of    intertrigo    set    up    by 

Oxyuris  vermicularis ,   696 
Microgametes  of  Coccidiidea,   137,   139,   140 
Microscope,  use  of,  discoveries  of  parasites 

from,  10 
Microsporidia,   129,   184 

—  characters  and  habitat,  28 

—  morphology  of,   185 

—  various  pathogenic  members,  186 
Midges,  see   Chironomidce,   Ceratopogoninoe, 

Psychodidce 

Miescher's  tubes,   187,   188 

Mikrofilaria  hancrofti,  prevalence  in  blood, 
prevalence  of  filarial  diseases  propor- 
tionate to,,  400 

and  Mikroloa  lea,  distinction  be- 
tween, 398 

—  diurna,  larvae  of  Loa  loa,  412 
presence  in  blood,  412,  414 

—  perstans,  morphology,  416 

and  M.  diurna,  simultaneous  presence 

in  blood,  414 

—  philippinensis ,  407 

—  powelli,    407 
Microfilariae,  periodic,  393,  394 

Milk    cure    in    expulsion    of    Strongyloides 

stercoralis,  675 
Milton,  bilharzial  vaginitis,  643 

—  treatment  of  bilharziasis,  643 
Mimomvia,  characters,  565 


INDEX 


873 


Miiicliiii  on  genus  Entamceha,  733 

—  researches  on   Trypanosoma  lewisi,  89-92 

—  see  also  Nicoll  and  Minchin 

Mineliin   and  Fantham,  on  Rhinosporidium 

kinealyi,  195,  196,   197 
Minchin    and    Woodcock    on    Trypanosoma 

noctuoB,   737 
Mineral  waters  in  intestinal  myiasis,  728 
Miners,  prophylaxis  against  ancylostomiasis 

in,  684 
Mines  infected  with  ancylostomes,  disinfec- 
tion of,  685 
Miracidia    of    digenetic    trematodes,    mor- 
phology of,  226,  227 
Miracidium,   germ   cells  of,   227 
Mitchell,  treatment  of  Oriental  sore,  628 
Mites    attacking    man,    geographical    distri- 
bution of  species,  486 

—  case    of    so-called    dysentery    said    to    be 

due  to,  512 

—  living  endoparasitically  in  animals  and 

birds,  491 

—  see  also  Acarina 

—  see  also  Arachnoidea 
Mochlonyx,  565 

Moiriez,    species    of    Chorioptes    found    on 

man,   521 
Moldovan,     schizogony     in     Leucocytozoon 

ziemanni,  153 
Molluscs,  spirochsetes  in,  114 

—  fresh-water,     round    Cairo,    cercariae    of 

bilharzia  type   in,  277 
Molluscum  contagiosum,  207,  208 
Monas  pyophila,  62 

characters  of,  62 

Mondiere,      perforation     of     appendix     by 

Ascaris,  655 
Monera,  26 
Moniez,    Aleurobius    (Tyroglyphus)   farinoB, 

511 

—  on  derivation  of  entozoa,  21 
Monkeys,     dysentery     in,     associated    with 

presence      of      (Esophagostomum     apio- 
stomum,  444 

—  experimental    infection   with   Spirochmta 

duttoni,  117 

—  inoculation  experiments  with  yaws  upon, 

128 

—  rabid,  spinal  ganglia  of,  cultivation,  210 

—  Trypanosoma  simicB  virulent  to,  100 
Monocystis,  hosts  of,  134,  135 

—  agilis   from   seminal   vessels   of  Lumbri- 

cus,   130,   132 

life-cycle  of,  132,  133 

Monogenea,   canalis   vitello-intestinalis,   222 

—  ova  of,   deposition,   223,  224 

—  post-embryonic   development   in,   224 
Monostomum  lentis,  244 

Monothalamia  (testaceous  amoebae),  charac- 
ters of,  47 

Montgomery,  transmission  of  rinderpest.  742 
Moore    and    Breinl,    latent   bodies    of    Try- 
panosoma gainbiense,  77 
Moosbrugger,     earth-eating     in     connection 
with  Trichuris  trichiura  infection,  679 

—  trichocephalus   anaemia,   651 


Moriggia,    Glyciphagus  cursor,  513 
Morkowitin,    Oxyuris    infection   in   relation 

to  appendicitis,  653 
Morphia,  injection  of,  in  relief  of  griping 

and  straining  in  amoebic  dysentery,  618, 

619 
Morsasca,  trichocephalus  anaemia,  651 
Moscato,    chyluria    following    infection    by 

Eustrongylus   gigas,   682 
Mosquito  nets,  use  of,  636 

—  worm  in  Trinidad,  598 

how  destroyed,  598 

Mosquitoes,   abdomen,   550 

—  acting  as  hosts  of  Filaria  bancrofti,  398 

—  alimentary  canal,  550,  551 

—  anatomical  remarks  on,  548 

—  antennae  of,  548 

—  aquatic  in  larval  and  pupal  stages,  555 

—  breeding  places  of,  553,  557 

—  campaign     against,     in     prevention     of 

malaria,  636 

—  copulation  of,  553 

—  distinguishing   features   of   Chironomidce 

(midges)   from,   579 

—  females  alone  blood-suckers,  552 

fertilized  in  autumn,  hibernation  of, 

555 

—  first  development  of  malarial  parasite  in, 

traced  in  Plasmodium  relictum,  170 

—  labrum,  labium,  and  hypopharynx,  548, 

549 

—  larvae,  food  of,  557 

living  in   salt  water,  557 

position  assumed  in  water,  557 

—  length  of  egg,  larval  and  pupal  life,  555 

—  maxillae  and  mandibles,  548,  549 

—  ova  of,  558 

float   on   water,   559 

— •  —  localities  for  deposition  of,  553 

—  proboscis  of,  548 

—  pupae  of,  558 

—  spread  of  malaria  in  birds  by,  158 

—  systematic  remarks  on,  548 

—  typical   structure   of,    diagram   showing, 

558 

—  ubiquitous  existence  of,  555 

—  see  also  Culicidm 

Moth-like  appearance  of  PsychodidcB,  581 
Mott,  F.  W.,  association  of  Treponema  with 

general   paralysis,   125 
Moty,     Oxyuris    infection     in     relation    to 

appendicitis,  653 
Mouqui,  mite  attacking  man,  486 
Mouth,    human,    cultivation    of    species    of 

Treponema  from,  741 

spirochsetes  in,   122,   740 

— •  infection      with      Oxyuris      vermicularis 

solely  through,  469 

—  maggots  in,  721 
Mucidus,  characters,  563,  571 

Mulder,    infection    with    Demodex    follicu- 

lorum,  708 
Mules,  murrina  in,  trypanosome  causing,  98 

—  "surra"  in,  95 

Miiller,     D.,     echinococcus     cysts     causing 
urticaria,  651 


874 


THE    ANIMAL    PARASITES    OF    MAN 


Miiller,   J.,   discovery   of  Myxosporidia,   181 

—  O.    F.,    discovery    of    and    views    as    to 

cercarise,   12 

of  origin  of  tapeworms  by,   11 

Miiller's  psorosperms,  135 

Murrina  in  mules,  trypanosome  causing",  98 

Musca  domestica  (common  house-fly),  586 

breeding  grounds,  destruction  of,  586 

characters,  585,  586 

destruction  of,   methods,   586 

diseases  spread  by,  586 

hibernation  as  puparia,  586 

larvae    (maggots)  of,  characters,   586 

life-cycle  of,  586 

ova  of,  places  where  deposited,  586 

pupa  of,  586 

—  pattoni,  611 
MuscidoB,  584 

—  African,  larvae  of,  590 

causing  myiasis  in  man  (footnote), 

590 

—  blood-sucking,  603 

—  larvae  of,  other  than  Lucilia,  in  nose,  720 
Muscles,  encystment  of  Trichinella  spiralis 

in,  425 

—  invasion  by  Trichinella  spiralis,  424,  425 

—  of  nematodes,  361 

—  sarcosporidia  in,   191 
Muscular  system  of  Hiriidinea,  480 
Musgrave,  on  human  intestinal  amoebae,  31 

—  and   Clegg's   culture   media   for   amoebae, 

743 
Mussels,  fresh-water,  spirochaetes  of,  114 
Mutualists,  nature  of,  6 
Myiasis,  715 

—  auricular,   615 
treatment,   615 

—  dermatosa  oestrosa,  725 

—  due  to  Sarcophaga,  589,  590 

—  externa,  715 

methods    of    treatment    recommended 

by  various   authors,   719,   720 
rare  situations  of,  723 

—  gastric,  treatment,   728 

—  human,  occurring  in  mountains  of  Cen- 

tral Sahara,  598 

—  in  French  West  Africa,  cause  of,  614 

—  intestinal,  725,  726 
chronic,   726 

complicated  by  mucous  colitis,  726, 

727 

diagnosis,   728 

irrigation  of  rectum  in,  728 

larvae  of  different  species  of  flies  found    I 

in,  728  ! 

modes  of  infection,  727  i 

views  of  various  authors  on,  727 

prognosis,   728 

prophylaxis,  728 

symptoms,  726 

treatment,  728 

and  cutaneous,  fly  causing,  585 

—  larvae     of     African      Muscidoe      causing 

(footnote),   590 

—  nasal,  715 

cases  of,   authors  reporting,  716,  717 


Myiasis,  nasal,  connection  with  ozaena,  717, 
722,  723 

discharge   from  nose  in,  718 

due   to   Sarcophaga,   treatment,   723 

fatal  termination  of,  718 

from  Sarcophaga  wohlfahrti,  722,  723 

maggots  of  flies  setting  up,  588 

prophylaxis   against,   718 

symptoms,  717,  718 

treatment,  719 

see  also  Peenash 

—  cestrosa,  geographical  distribution,  724 

prevalent  among  rural  population,  724 

rare  in  man,  724 

treatment,  725 

Myriapoda  parasitic  in   intestine   and   nose 

of  man,   483 
MyxidiidcB,  184 
Myxidium  lieberkiihni,  182 
Myxobolidce,   184 
Myxobolus  cyprini,   184 

—  pfeifferi,  184 

— cause  of  barbel  disease,  184 

spore  formation,   183 

—  neurohius,  184 

—  schematic  representation  of  spore  of,  182 
Myxoedematous  form  of  Brazilian  trypano- 
somiasis, 88 

Myxosporidia,  129,  181 

—  authors  adding  to  knowledge  of,  182 
describing  species  causing  diseases  in 

fishes,   182 

—  characters  and  habitat,  28,   182 

—  free  forms  of,  182 

—  introduction  of  term  of,  by  Biitschli,  181 

—  invasion    by,    causing    disease    in    fishes, 

182,  184 

—  mode  of  infection,   184 

—  multinucleate  trophozoite  of,  182 

—  plasmotomy,    182 

—  spore  formation,  182,  183 

—  tissue  parasites,   182 
Myzomyia,  characters,  561,  567 

—  funesta,  breeding  places  of,  557 
Myzorhynchella,  characters,  561,  568 
Myzorhynchus,  characters,  562,  568 


Nabarro,  on  sleeping  sickness,  68 

—  on   Spirochmta   duttoni,   116 

Nagana   (tsetse-fly  disease),  agent  of  trans- 
mission, 93 

—  fatal  to  horses,  asses  and  dogs,  94 

—  prevalent  among  and  generally  fatal  to 

cattle,  93,  94 
---  treatment  by  arsenic,  94 

—  trypanosomes  in  blood  of  horses   suffer- 

ing from,  68 
Nagel,   chloroform   and   syrup   of   senna  in 
expulsion  of  ancylostomes,   686 

—  filmaron    in    expulsion    of    ancylostomes, 

687 
Nagelschmidt,  treatment  of  scabies,  706 


INDEX 


875 


Naphthalene  iu  evacuation  of  Oxyuridce,  697 

—  in  intestinal  myiasis,  728 

Naphthol    ointment,    dressings    of,    in    head 

louse  infection,  710 

(Kaposi's),  application  in  scabies,  707 

Nasal  cavity,  deposition  of  ova  of  Oestrus 

ovis   in,   598 

see  also  Myiasis,  nasal 

Ling-uatula  parasitic  in,  523,  524,  526, 

527 

—  polypus,    Bhinosporidium    causing,    195- 

197 

Nason,  Ascaris  in  appendix,  causing  in- 
testinal  obstruction,   654 

Nasse,  investigations  of  Coccidia,  135 

Natal,  larva  of,  characters,  591 

Nattan-Larrier,  cross-immunity  experiments 
with  trypanosomes,  80 

—  Tetramitus  mesnili,   57,  624 

Natural  flagellates  of  insects,  103,  104,  107, 

112,  739 
Naunyn,    mode    of    formation    of    daughter 

cysts   of   echinococcus,    352 
Neave,    S.,    ulcers    set    up    by    invasion    by 

larvae  of  Cordylobia  anthropophaga,  592 
Necator,  447 

—  americanus,  450 

geographical  distribution,  459 

habitat,  459 

—  —  morphology,  457,  458 

organs  compared  with  those  of  Ancylo- 

stoma  duodenale,  458 

—  characters,  457 

—  exilidens,  characters,  459 
habitat,  459 

Negri,  experimental  infection  with  Sarco- 
cystis  muris,  192 

—  on   Neuroryctes,   208 
Negri's  bodies,  208,  209 

Neligan,  Leishmania  tropica  in  dogs,  108 

Nemathelminthes,   360 

Nematoda,   see   Nematodes 

Nematode  larvae  in  blood  in  cases  of  pru- 
ritus, 378 

in   periosteum  of  upper  jaw   in   case 

of  gingivitis,  378 

Nematodes,   anatomy  of,  360 

—  bursa  copulatrix  of  males,  370 

—  chorion  enveloping  ova,  371 

—  classification  of,  374 

—  clearing   of,   473 

—  cutaneous  glands,  unicellular,  361 

—  cuticle  of,  360 

—  cutis   of,   361 

—  dermo-muscular    layer    of,    361 

—  development  of,  17,  371  ' 

—  embryos,  372 

—  encapsuled  forms  of,  17 

—  epithelium  of,  360 

—  excretory  canals,  anterior,  367 
organs,  366,   367 

special,  lacking  in  certain  genera, 

367 

—  —  pore  and  duct,  367 
vesicle,  367 

—  fixation  of,  473 

—  oflandular  stomach  of,  363 


Nematodes,   gubernaculum   of  male   genital 
apparatus,  369 

—  hatched   from   eggs  of  Sphcerularia,   5 

—  heterogony   in,   372 

—  hind  gut,  363 

— ■  infection   by,   644 

—  intestinal  caeca,  364 
canal,  363 

—  "isolation  tissue,"  362,  363,  364 

—  life  spent  in  intermediate  and  final  host, 

18 

—  marine,  ventral  gland  of  (so-called),  367 

—  mounting  head  of,  473 

—  muscles  of,  361 

—  nervous  system,   364-366 

—  observed  in  human  eye,  412 
in  man,  376 

—  oesophageal  glands,  364 

—  oesophagus   of,   363 

—  organs    of    sense     lacking    in     parasitic 

species,  366 

—  ova  of,  371 

conveyance  to  definite  host  with  inter- 
mediate  host,  373 

without  intermediate  host,  372 

detection,  473 

developmental  capacity,  371,  372 

—  ovejector,  368 

—  oviduct,  368 

—  parasitic  and  free-living,   connection,  20 

—  preservation  and  examination  of,  473 

—  rolling  of,  473 

—  seminal  receptacle,  368 

—  sexual  organs,   367 

female,  367,  368 

diagram   of,   368 

male,  369 

diagram  of,  368 

—  small,  detection  of,  473 

—  spicules  of  male  genital  apparatus,  369 

—  staining  of,  473 

—  testis  of,  369 

—  "  tuft-like  "  or  "  phagocytic  "  organs,  362 

—  viviparous  species,  371 

—  young,  skin  diseases  due  to,  in  dogs,  378 
Nematodirus,   habitat,   438 

—  morphology,  438 
Neocellia,  characters,  562,  569 
Neomyzomyia,   characters,   561,  567 
Neopsylla,  distinctive  characters,  545 
Neosalvarsan  in  syphilis,  632 

—  in  yaws,  632 
Neosporidia,  129,   181 

—  characters,    28,    129,    181 
Nephrophages  sanguinarius,  characters  and 

morphology,  490 

presence   in   urine,   490 

Nervous  system  of  Cestoda,  289,  290 

of  Echinorhynchus,  475 

of   Hirudinea,   481 

of  Insecta,  530 

of  nematodes,  364-366 

central,  effect  of  Dipylidium  caninum 

on,  649 

Neumann,  mosquitoes  transmitting  Plasmo- 
dium relictum,  170 


876 


THE   ANIMAL    PARASITES    OF    MAN 


Neumann,     podophyllin     in      expulsion     of 
ancylostomes,  687 

—  synopsis  of  genus  Ornithodorus,  508 

—  jbable  of  species  of  Argas,  505 
Neuritis,  optic,  following  male  fern  poison- 
ing, 670 

Neuroptera,  characters,  531 
Neuroryctes,  208 

—  hydrophobioe,  208 

minute   granules   in,   210 

Neurosporidium,   195 

—  cephalodisci,  195 

Newstead,  Amblyomma  cayennense,  501 

—  life-cycle  of  Phlebotomus,  582 

■ —  Margaropus  anmUatus  aiistralis,  505 

—  means  of  separating  sjDecies  of  Glossina, 

604 
Niaibi,  mite  attacking  man,  486 
Nicoll,     development    of    cestodes    without 

intermediate  host,   17 
■ —  and   Minchin,   cysticercoids   in   rat  fleas, 

327,    328 
Nicolle,  immunity  experiments  with  Leish- 

mania  infantum  and  L.  tropica,  112 

—  and    others,    transmission    of    relapsing 

fever  by  lice,   120,   121 
Nicollia,  174 

—  quadrigemina,  174 

Nicotiana  soap,   application  in   scabies,  707 
Nits,  methods  of  getting  rid  of,  from  hair, 

710 
Nitzsch,  views  as  to  cercariae,   12 
Noc,  cultivation  of  species  of  amoeba  by,  41 

—  on  Lamblia  intestinalis ,  60,  625 
Noguchi,  cultivation  of  parasite  of  rabies,  210 
of  Treponema  from  human  mouth, 

128,   741 
of  Treponema  pallidum,  125 

—  method  of  cultivation  of  spirochsetes,  123 

—  Spirochoeta  phagedenis,  122 

—  Treponema  calligyrum,   126 

—  and     Cohen,     cultivation     of     so-called 

trachoma  bodies,  210 

—  and    Moore,    association    of    Treponema 

with  general  paralysis,  125 
Noller,  development  of  Trypanosoma  lewisi 
in  dog  flea  (Ctenocephalus  canis),  90,  92 

—  method   of    controlling    fleas    during    ex- 

periments, 93 

Nordmann,  von,  discovery  of  miracidia  of 
flukes,    12 

Normand,  association  of  amoebae  with 
colitis,  30 

of  Strongyloides  stercoralis  with  diar- 
rhoea, 380 

Norway  itch   (scabies  norvegica),  520 

Nose,  ascarides  in,  690 

—  dipterous    larvae    in,    in    enormous    num- 

bers, 716,  717 

—  discharge  from,  in  nasal  myiasis,   718 

—  human,  Myriapoda  parasitic  in,  483 

—  larvae  of  Hypoderma  bovis  in,  724 

of  Lucilia  macellaria  in,  715,  716 

see  also  Myiasis,  nasal 

of  Oxyuris  vermicularis  in,  469 

—  leeches  in,  700,  701 


Nose,   leeches   in,   causing   epistaxis,   701 

—  maggots  in,  588 

—  Oxyiiridoi  migrating  into,  695,  696 
Nosema  apis,  184 

- —  —  life-cycle  of,   185 

pansporoblast  and  sporoblast    >f,  185 

planont  of,   185 

—  bombycis,  184 

spores  of,  186 

Notoedres  cati,  521 

—  cuniculi,  521 

—  notoedres,  521 

Novy  and  MacNeal,  artificial  cultivation  of 
trypanosomes,  69 

Novy -MacNeal  medium,  744 

Mathis's  modification,  744 

Novy-MacNeal-Nicolle  medium,  best  for  cul- 
tivation of  Leishmania  infantum,  109 

for  cultivation  of  Leishmania  tropica, 

108 

formula,  744 

Nut-pickers  affected  by  Leptus  aiitumnalis 
(footnote),  485 

Nuttall,  Spirochoita  marchouxi,   119 

—  Piroplasmidce,    174 

Nuttall  and  Hadwen,  trypan-blue  in  treat- 
ment of  piroplasmosis,   178 

—  and      others,      nuclear      phenomena      of 

Babesia  canis,  176 

Theileria  parva,   179 

Nuttallia,  characters,   174 

—  equi,  cause  of  equine  piroplasmosis,  174, 

178 
life-cycle  in  red  blood  corpuscles,  173 

—  herpestidis ,   174 
Nycteribiidce,  611 
Nyctotherus,   204 
Nyctotherus  africanus,  206 

—  faba,  205 

—  —  morphology,  205 

—  giganteus,  205,  206 

morphology,  205 

Nyssorhynchus,  characters,  562,  569 

0. 

Occiput,  abscess  of,   liver-fluke  in,  243 
Ochindundu,  bite  of,  541 

—  characters   of,   541 

Ochromyia  anthropophaga ,  larvae  of,  charac- 
ters  (footnote),  590,  591 

—  —  —  hosts  of,  590 

(Edema  following  bite  of  Argas  reflexus,o06 
Oerley,  induction  of  facultative  parasitism 

of  Bhabditis  pellio,  377 
(Esophageal  glands  of  nematodes,  364 
CEsophagostomeoe,  characters,  439 
(Esophagostomum,  morphology,  441 

—  apiostomum,  habitat  and  host  of,  444 
morphology,   444 

—  brumpti,  habitat,  441 
morphology,  441 

—  stephanostomum,  habitat,  444 

var.    thomasi,    morphology,    442,    443, 

444 
(Esophagus  of  Hirudinea,  480 

—  of   nematodes,   363 


INDEX 


877 


CEsophagus,  trichomonads  in,  00 
Oestrijoe   (wabble   flies),   boils  produced  by, 
725 

—  cavicolous,  598 

—  cutaneous,   595 

—  flight  time  of,  725 

—  gastricolous,   599 

infection   by,   729 

see  also  Creeping  disease 

—  hosts   of,   594 

—  larvae  of,  occurrence  in  man  rare,  724 

—  method    of    depositing    ova    on    skin    of 

man,   725 
Oestrus   {Cephalomyia)  oris,  598 

geographical   distribution,   598 

ova  of,  deposition  in  nasal  cavity, 

598 
Oil,  injections  of,  in  nasal  myiasis,  719 
Ointment,  application  in  scabies,  706 
Oken,  views  as  to  origin  of  cercariae,  12 
Oleum  chenopodii  in  ancylostomiasis,  754 
Oligosporulea,  195 
Oligotricha,  29 
Oliver,  artificial  infection  of  human  beings 

M^ith   Cysticercus   bovis,   340 
Omentum,   abscess  of,   with  Ascaris   ova  in 

pus,  657 
Onii,   diagnostic   sign,  of   presence   of   Spar- 

ganum  mansoni  in  body,  659 
Onchocerca,  417 

—  volvulus,    417 

—  —  distribution  in  West  Africa,  419 

— -  —  invading  lymphatic  vessels,  418,  419 

invasion     in     man      associated     with 

formation  of  tumours,  418 

—  —  measurements,  755 

morphology,   417,   418 

Onchocercinw,  417 

Oncospheres  (embryos)  of  tapeworms,  298, 
299 

certain  species  of  animals  neces- 
sary for,  299 

development  into  plerocercoid,  300 

further     develojjment     must     take 

place  in  suitable  animals,  299 

—  migration   in   body,   302 
O'Neil,  filaria  infection  of  skin,  378 
Onions,  Anguillulina  putrefaciens  living  in, 

379 
Oocysts  of  Cuccidiidea,   141 

—  of   malarial   parasites,    163 
Opalina,   198,  207 

—  ranarum,  207 
Ophryocystis,  135 

Op  isthorchiidcB ,  morphology,  232 
Opisthorchiinw ,  252 
Opisthorchis,  252 

—  felineus,  development,  254 

—  —  geographical  distribution,  252 
hosts  of,  252 

intermediate,   254 

mode  of  infection  by,  254 

morphology,   252 

synonyms,  252 

—  pseudo  felineus,     anatomy     of,     diagram 

illustrating,  254 


Opisthorchis   sp.,   habitat,   753 

morphology,   753 

Opisthotonos,  disappearance  after  exjiulsion 

of  Ascaridce,  649 
Oppenheim,  maculae  caerulae   (taches   bleues) 

due  to  infection  by  crab  louse,  7l2 

—  treatment  of  crab  louse  infection,  712 
Oppila^ao,  synonym  of  Brazilian  trypaiioso- 

miasis,  87 
Oral   cavity,   cancer  of,  association   of  Ent- 
amoeba buccalis  with,  43 

trichomonads   in,   55,   56 

Orbit,    cysticercus   of,    664 
Orbital  cavity,   Pycnosoma  maggots   invad- 
ing, 588 
Orchitis    from    Filaria    bancrojti    infection, 

677 
Oribates  sp.,  489 
Oriental  sore,  cause  of,   107,  627 

experimental  production,   109 

geographical  distribution,   108 

germ  of,  possible  carrier,  580 

immunity  to,  procured  by  inocula- 
tion, 108 

objection  to  name,  107 

occurrence  in  dogs,   108 

parasite  producing,   107,  627 

pathology  of,  627 

—  —  preventive  measures,  628 

relation     of    infantile     kala-azar    to, 

shown   experimentally,   109 

sites  of  occurrence  on  body,  108 

transmission  of,  bugs  possibly  con- 
nected  with,    108,   536 

treatment,   628 

Ornithodorus  coriaceus,  geographical  distri- 
bution, 509 

—  megnini,  characters,  510 

ears  of  hosts  infested  by,  510 

geographical  distribution,  510 

hosts  of,  510 

—  moubata,  carrier  of  African  tick  fever^ 

116,  496 

of  Filaria  perstans,  508 

of   spirochaete    of   relapsing    fevei*, 

508 

geographical  distribution,  509 

immunity      against      infection      with 

Spirochceta  duttoni,  119 

length  of  life  apart  from  host,  495 

Malpighian  secretion  passed  by,  sig- 
nificance, 117 

transmission    of    Spirochceta    duttoni 

by,  116 

—  savignyi,  509 

geographical  distribution,  509 

transmitting  Spirochceta  duttoni,  739 

—  synopsis  of  genus,  508 

—  talaje,   119 
bite  of,  509 

geographical  distribution,  509 

—  tholozani,  geographical  distribution,  51Q 

—  turicata,   119 

bite  of,  effects,  509 

Omithomyia  lagopodis,  bite  of,  611 
Orthoptera,   characters,   531 


878 


THE   ANIMAL    PARASITES    OF    MAN 


Otter,  Brazilian,  liost  of  Paragonimus  rudis, 

251 
Ova,  transmission  of  intestinal  worms  by,  11 

—  see  also  under  Names  of  parasites 
Owen,  Trichina  spiralis,  423 

Ox,  liver  of,  Echinococeus  multilocularis  in, 
357 

—  Sarcocystis   hlanchardi  from,   190 

—  gad  fly  (Tabanus  hovinus),  601 
Oxazine     producing    blepharoplastless    try- 

panosomes,    101 
Oxen,  amount  of  prevalence  of  Cysticercus 
bovis  in,  340,  341 

—  echinococci   in,   346 

—  how     infected     with     Para7nphistomum 

cervi,   226 
Oxygen   necessary   in    cultivation   of   spiro- 

chaetes,   123 
Oxyuriasis,   diagnosis,   696 

—  dysentery  followed  by  recovery  from,  698 

—  in  children,  695 
treatment,   697,   698 

—  prophylaxis  against,  697 

—  treatment  by  drugs  and  purgatives,  697 
local,  697 

Oxyuridce,  467 

—  migrating  into  nose,  695,  696 

—  morphology,  375 

—  relationship  to  appendicitis,  698 
Oxyuris,  467 

—  ambigua,  469 

—  compar,  469 

—  curvula,  469 

—  encapsuled  in  female  pelvis,  657 

—  in  appendix,  654,  655 

—  infection  in  relation  to  appendicitis,  653 

—  invading   peritoneal   cavity,   657 

—  lacks   intermediate   host,   21 

—  mastigodes,  469 

—  poculum,  469 

—  tenuicauda,  469 

—  toxic  action  of,  651 

vermicularis,    association    with    appendi- 
citis and  typhlitis,  467 

development,  468 

direct,  469 

experimental  self-infection  with,  469 

habitat,  467 

infection  by,  694 

with,  mode  of,  469 

.  —  larvae  of,  found  in  nose,  469 

life-history   of,   467,   468,   469 

males  rarely -met  with  in  faeces,  468 

migration  from  intestine,  lesions  and 

symptoms  of  irritation  set  up  by,  694, 
695 

of,   in    and   from   intestinal   tract, 

467 

morphology,  467 

ova  of,  where  deposited,  467 

supposed  origin  of,  11 

Oyster,   spirochaete  of,   114 
Ozaena,    connection    of   nasal   myiasis   with, 
717,    722,   723 


P. 

Page,    case    of    escape,    of    ascarides    from 

abdominal  operation  wound,  654,  655 
Paget,    observation    of    encapsuled    Trichi- 

nellae,  423 
Pallas,  on  transmission  of  intestinal  worms, 

11 
Panama,    larvicide    used    at,    in    campaign 

against  mosquitoes,  636 

—  Canal,     Stegomyia    fasciata     source     of 

danger   to,    574 

Pani-ghao,  skin  affection  set  up  by  pene- 
tration of  larvae  of  Ancylostomum  duo- 
denale,  455 

Panoplites,  577 

Pansporoblast,    183,   186 

Papataci  fever,  carrying  agent  of,  582 

Pappenheim's  panchrome  mixture,  751 

Paraboloid   condenser,   747 

Paraffin,  embedding  in,  for  sectioning  tissue 
parasitized  by  protozoa,  749 

Paragonimiasis,    639 

—  affecting  regions  other  than  lung,  639 

—  prophylaxis,  640 

—  see  also  Lung-fluke  disease 
Paragonimus,  morphology,  249 

—  compactus,  host  of,  251 

—  kellicotti,   hosts  of,  250 
spines  of,  251 

—  ringeri  (lung-fluke),  639 

development,  251 

diseases  caused  by,  251 

habitat,  251 

internal  organs,  diagram  illustrating, 

250 

morphology,  249,  250 

sites    of    body    in    which    found    post 

■mortem,   639 

spines  of,  251 

synonyms,   249 

—  rudis,  host  of,  251 

—  westermannii,  host  of,  250 

—  —  morphology,  diagram  illustrating,  250 
spines  of,  251 

Paraguay,    supposed   mode   of   transmission 
of  dermo-mucosal  leishmaniasis  in,  739 
Paralysis  due  to  tick  bites,  613 
geographical  distribution,  613 

—  of  dourine,   97 
Paramoeba,   44 

—  hominis,  45,  734 
characters  of,  45 

now  called  Craigia  hominis,  45,  734 

ParamphistomidoB,  231,  234 

ParamphistomincB,  231 

Paramphistomum  cervi,  method  of  infection 

of  oxen  by,  226 
Paraplasma,  180 

—  doubt  as  to  organismal  nature,   180 

—  occurs  naturally  in  guinea-pigs,  180 

—  flavigenum  possibly  associated  with  yel- 

low  fever,    180 
morphology,  180 

—  subflavigenum,  180 
Parasites,  definition,  1 


INDEX 


87$ 


Parasites,  derivation  of,  19 

—  diagnosis  of  presence  of,  10 

—  discoveries  from  use  of  microscope,  10 

—  great  fertility  of,  5 

—  hereditary  transmission  of,  19 

—  human,    Opisthorchis   felineus   most   fre- 

quently  found   at   autopsies   at   Tomsk, 
253 

—  incidental,  6 
human,  7 

—  influence  on  host,  8 

—  invading  many  hosts,  6 

—  limited  to  closely  related  hosts,  6 
to  one  species  of  host,  6 

—  migrations  in  host,  injuries  set  up  by,  9 

—  movements  of,   disorders  set  up  by,  9 

—  occasional   (temporary),  1 

—  origin  of,   10 

—  permanent,  bodily  changes  in,  3 

clasping  and  clinging  organs  in,  4 

classes  of,   2 

hermaphroditism  in,  4 

loss   of   organs   in,    3 

(stationary),  1,  2 

—  transference  from  one  host  to  another,  7 
Parasitic  life,  advantages  of,  20 
Parasitism,   facultative,   7 

Pariah  dogs,  liver  of,  habitat  of  Paropis- 
thorchis  caninus,  257 

North-west  Provinces,  India,  percent- 
age infected  with  Paropisth orchis  cani- 
nus,  257 

Paropisthorchis,  255 

—  caninus,  genital  pore,  255 
habitat,  257 

morphology,  255 

seminal  vesicle,   257 

synonyms,  255 

uterine  coils,  257 

vitellaria,  255 

Partridges,   Plasmodium    relictum  cause   of 

fatal    disease    in,    170 
Pasquale,   see  Kruse   and  Pasquale 
Pasteur,  L.,  researches  on  silkworm  disease, 

184 
Patterson,  maggots  of  Pycnosoma  removed 

from  orbital  cavity,  588 
Patton,   genus   Crithidia,   104 

—  Herpetomonas   muscce   doniesticce ,   102 

—  Piroplasma  gihsoni,   177 

—  places  Leishman-Donovan  body  in  genus 

Herpetomonas,   107 

—  probable     transmission     of     Leishmania, 

107,  108 

—  and     Cragg,     life-history     of     Lyperosia 

exigua,  610 
Peacock,  observation  of  encapsuled  Trichi- 

nellae,  423 
Pebrine    bodies    or    Nosema    bombycis    of 

Arthropoda,  184 
Pediculida>  (lice),  characters,  532 
Pediculoides  ventricosus,  effects  on  man,  469 

—  —  morphology,  489 

—  shape  of  preg-nant  female,  489 

—  —  synonyms,   489 

Pediculis  capitis  (head  louse),  characters 
and  morphology,  532,  533 


Pediculis_  capitis,  geographical  distribu- 
tion,   533 

habitat,  533 

infection  by,  709 

— causing  eczema,  709,  710 

diagnosis,  710 

greater  prevalence  among  females, 

709,  710 

remarkable  instances,  710 

resulting  in  blepharitis  and  con- 
junctivitis, 710 

in  plica  polonica,  710 

treatment,  710 

mouth  parts  of,  533 

ova  of,  533 

prophylaxis  against,  615,  616 

—  vestimenti  (clothes  louse),  characters,  533 
habitat,   533 

infection  by,  710 

lesions  and  symptoms  following,  711 

pest  among  soldiers  during  cam- 
paigns, 533 

prophylaxis  against,  615,  616 

transmission    of    relapsing    fever    by, 

120,  630 
Peenash  (nasal  myiasis),  588,  715 

—  ending  in  fatal  meningitis,  716 
Peiper,  cause  of  ancylostome  anaemia,  648 
Pelagutti,  treatment  of  cutaneous  and  mus- 
cular cysticerci,  663 

Pelletierinum  as  vermifuge,  673 

Pelvic  and  abdominal  organs,  blood-supply 
of,  as  illustrating  distribution  of  Schis- 
tosoma haematobium  in  body,  272 

Pelvis,  female,  Oxyuris  encapsuled  in,  657 

Pentastoma  armillatus,  hosts  of,  528 

—  denticulatum,    former    name    of    larval 

stage  of  Linguatula  rhinaria,  525,  526 

—  moniliformis,  hosts  and  habitat  of,  528 

synonyms   (footnote),  528 

PentastomidoB,  references  to,  528 
Pentateuch,  "  fiery  serpents  "  mentioned  in, 

probable  identification,  386 

Pentatrichomonas  bengalensis,  624,  735 

Pereira,  case  of  chorea  cured  after  expul- 
sion  of  Taenia,   648 

Perinseum,  tumours  of,  association  of  Dioc- 
tophyme  gigas  with,  431 

Peritoneal  cavity,  Ascaris  in,  656 

Oxyuris  invading,   657 

Peritonitis,  perforative,  due  to  Ascaris,  656 

Peritricha,  29,  200 

Perroncito,  artificial  infection  of  human 
beings  with  Cysticercus  bovis,  340 

—  infection  with  Lamblia  intestinalis ,  60 
Persia,    importation    of   African   tick    fever 

into,  613 
Persian    insect    powder    infusion    in    intes- 
tinal myiasis,  728 
Peru  oil,  application  in  scabies,  707 
Petrie,  treatment  of  bilharziasis,  643 
Petroleum  as  larvicide  in  campaign  against 
mosquitoes,  636 

—  dressings  of,  in  head  louse  infection,  710 

—  in   crab   louse   infection,   712 

—  and  benzine  in  crab  louse  infection,  712 


88o 


THE  ANIMAL   PARASITES    OF   MAN 


Pfeiffer,   L.,  pathogenicity  of  Coccidia,   136 
Pfeiffer,  R.,  Coccidia,  136 
Pharynx,  ascarides  invading,  691 

—  invasion     and     infection     in     man     by 

Fasciola  hepatica,  242 

—  leeches  in,  699 

—  of  Hirudinea,  480 
Philaematomyia,  position  of  genus,  611 

—  insignis,  611 

Philippine  Islands,  experiments  on  amoebae 

in,  618 
Philips,     eucalyptus     oil     in     expulsion     of 

ancylostomes,  687 
Phillips,   L.    P.,   on   Musgrave   and   Clegg's 

medium,  743 
treatment    of    balantidian    dysentery, 

637 
Phlehotomince,   characters,   581 
Phlebotomus,    blood-sucking,    581 

—  characters,  581 

—  duboscii,  582 

—  intermedins,  582 

—  geographical  distribution,  581 
of  species,  582 

—  larvae  of,  habitat,  582 

—  longipalpis,  582 

—  papatacii,  581,  582 

—  squamiventris ,  582 
Phonergates   hicoloripes,   541 
Phoniomyia,  characters,  565 
Phora  rufipes,  589 

larvae   (maggots)  of,  habitat,  583 

Phoridce,  characters,  582 

Phthiriasis,  agents  of,  533 

Phthirius  inguinalis  (crab  louse),  charac- 
ters, 534 

habitat,  534 

infection  by,  diagnosis,  712 

how  eifected,  711 

lesions    and    symptoms    following, 

711,  712 

sites  of  body  affected,  711 

treatment,  712 

rapid  reproduction  of,  534 

—  pubis,   prophylaxis   against,   616 
Phthisis,  filaria  associated  with,  408 
Physaloptera,  habitat  and  hosts  of  species, 

460 

—  caucasica,  morphology,  461 

—  mordens,  geographical  distribution,  461 
habitat  and  host,  461 

morphology,   461,   462 

—  morphology,  460 
Physalopteridce,  375,  460 
Phytoparasites,  1 

Pierantoni,  Agamofilaria  labialis,  407 
Pig   concerned   in    transmission   of   Balanti- 
dium  coli,  202 

—  development    of   Trichinella   spiralis   in, 

426,  427 

—  domestic,  normal  host  of  Cysticercus  cel- 

luloscp,   332 

—  echinococci  in,  346 

—  geoQ-raphical      distribution      of      Tmnia 

solium  corresponds  with  that  of,  334 

—  host  of  Paragonimus  kellicotti,  250 


Pig,  intestine  of,  Fasciolopsis  buski  in,  246 

—  Metastrongylus  apri  in,  433 

—  organs   infected   with   echinococcus,    per- 

centage of  frequency,  347 

—  rectum  of,  Balantidium  coli  present  in, 

202 

—  Sarcocystis  miescheriana  in,   190 

—  Sarcosporidia  in,   187 

—  trichinous,     proportion     to     healthy,     in 

Prussia,  429,  430 

—  Trypanosoma  simios  virulent  to,  100 
Pigeon  lofts  inhabited  by  Argas  rejlexus,  506 
Piophila  casei,  characters,  583 

larvae  of,  found  in  faeces,  583 

in  nose,  720 

Piroplasma,  172,  173,  174 
— •  see  Babesia 

—  gibsoni,   111 

—  hosts   of,    173,    174 
PiroplasmidoB,    172,   742 

—  genera  of,   174 
Piroplasmosis,  treatment  of,  178 

by  trypan-blue,  178 

symptoms  of,  178 

transmission  by  ticks  from  recovered 

to  uninfected  animals,   178 
Placobdella,  482 

—  catenigera,      geographical      distribution, 

482 
Plague,  fleas  carriers  of,  543,  547 
Planont,    185 
Plants,    flagellosis    of,    possible    connection 

with   leishmaniasis,   739 
Plasmodium,    151,    742 

—  falciparum,  see  Laverania  malarias 

—  malaricB,   development  in  red  corpuscles 

of  man,  asexual  stage,   166 

distinctive  characters,  167 

lesions  set  up  by,  not  marked,  634 

parasite  of  quartan  malaria,  166,  633 

pigment  granules  of,  166,  167 

schizogony  of,  166 

synonyms,  166 

trophozoites  of,   differ  from   those  of 

tertian   parasite,   166 

—  or    haemamoeba    type    of    Hoemosporidia 

includes  malarial  parasites  of  man  and 
birds,  151 

—  relictum,   first   development   of   malarial 

parasite  in  mosquito  traced  in,  170 

hosts  of,  170 

mosquitoes  transmitting,   170 

stages  in  life-history,   170 

synonyms,  170 

—  species,  differential  table  of,  171 

—  tenue,  170 

—  vivax,  agent  of  simple  tertian  malarial 

fever,    164,    633 
cultivation  of,  clumping  not  observed 

in,  172 
number  of  spores  produced,   172 

—  —  development   in   red   blood   corpuscles 

of  man,  160,  164,  165 

—  —  —  of  ''Polymitus,"    160,   165 

time  occupied  by,   165 

distinctive  characters  of,  166 


INDEX 


88l 


Plasniociium  rivax,  lesions  set  up  by,  uot 
marked,   634 

life-cycle   of,    160,    164 

merozoites   of,    165 

migration,    165 

—  —  micro-  and  macrogametocytes  of,  165 
—  23igment  granules,  165 

small  variety,   166 

—  —  •'  stippling,"    165 

synonyms,  164 

Platylielminthes  (flat  worms),  211 

—  central    nervous   system   of,    211 

—  classification,   212 
— -  definition,    211 

—  diseases  caused  by,  638 

—  excretory  apparatus,  211 

—  hermapliroditic,  211 

—  integument   of   body    of,    211 

—  method  of  reproduction,  211 

—  morphology,  211 
Plerocercoid,   definition  of,  301 
Plerocercus,  definition  of,  301 

Pleurae,  invasion  by  Faragonimus  ringeri, 
251 

Plica  polonica  due  to  head  louse  infection, 
709,  710 

treatment,  710 

Plimmer,  H.  G.,  treatment  of  sleeping  sick- 
ness with  antimony,   623 

and  Bradford,  Sir  J.  Eose,  Trypano- 
soma brucei,  93 

Pliny,   Ascaris   lumhricoides   known   to,  464 

Pneumocystis  carinii,  90 

Pneumocysts  in  rats,  90 

Pocock,  geographical  distribution  of  Orni- 
thodorus  mouhata,  508,  509 

Podophyllin  in  expulsion  of  ancylostomes, 
687 

Polar  capsule,  181,  183,  184,  186 

—  filament,  183,  184,  186 

Polecat,    intestine    of,     Isospora    higernina 

parasitic  in,   149 
Poliomyelitis  acuta,  possible  rhizopods  in,  46 

—  carrier  of,   610 

—  epidemic,   insects   transmitting,   612 

—  virus  of,  536 

carried  by  house-fly,  586 

Pollack,   invasion   by  Loa   loa,   678 

Polymastigina,  52 

Polymitus  of  Plasmodium  vivax,  160,  165 

—  form  of  malarial  parasites  (footnote),  162 
Polypus,     nasal,     caused     by     Rhinospori- 

dium-    kinealyi,    195,    196 
Polysporea,  182,  184 
Polysporulea,   195 

Polystoynum  integerrimum,  organs  of,  218 
Ponds,  mosquitoes  depositing  ova  in,  553 
Pork,  cysticerci  in,  cause  of  decrease,  334 

—  eating  of,  cause  of  trichinosis,  423 
means    of   infecting   man   with   cysti- 
cerci, 334 

—  inspection    of,    in    prophylaxis    against 

trichinosis,  429 
Porocephalus,  523 

—  armillatus ,  527 

synonyms  (footnote),  528 


Porocephalus   constrictus,   characters,   526 

hosts  of,   526,   527 

organs  of  body  invaded  by,   526,  527 

synonyms,  526 

Port  Natal  sickness  (Cape  ailment),  488 

Portal  vein  and  liver.  Schistosoma  hcema- 
tobium  most  easily  found  post  mortem 
in,  273 

and  vena  cava,  communication  be- 
tween, how  formed,  272 

tributaries  of,  as  illustrating  distri- 
bution of  Schistosoma  hcematobium  in 
body,  272 

Porter,   A.,   Crithidia  pulicis.    111 

generic      differences      among      insect 

flagellates,    103    (fig.    49) 

Herpetomonas   muscm   domesticre,   102 

Leucocytogregarina,  154 

Theileria  parva,  179 

—  see  also  Fantham  and  Porter 
Portschinsky,   deposition   of  ova  of   Oestrus 

oris,  598 

—  method  of  destroying   Tabanidce,   601 
Posner,  case  of  amoebae  in  urine,  46 
Posselt,    cutaneous    tumours    due    to    cysti- 
cerci, 662 

—  reasons    for    distinction    of    multilocular 

from     hydatid     or     unilocular     echino- 

coccus,   358 
Post-flagellate  stage  in   herpetomonads,   103 

in   Crithidia,   104 

Potassium  iodide  in  treatment  of  cutaneous 

and  muscular   cysticerci,   663 

—  permanganate,    application    in    Oriental 

sore,   628 
Pou  d'agouti,  mite  attacking  man,  486 
Poultry,    fatal    epizootics    among,    due    to 

Eimeria  avium,  142 
Poultrymen     attacked     with     Dermanyssus 

gallince,  493 
Poupee-Desportes,    Guinea    worm    infection, 

676 
Powell,    method    of    destruction    of    Sarco- 

phaga  larvae,   723 
Predtetschensky,  expulsion  of  Hymenolepis 

nana,  661,  662 
Pre-flagellate  stage  in  herpetomonads,   103 

in   Crithidia,   104 

Price,     Dodds,     method     of     prevention     of 

Indian  kala-azar,  627,  739 
Prima,  fatal  case  of  myiasis  externa,  716 
Privies,     disinfection    of,     as     prophylactic 

against  ancylostomiasis,  685 
Proflagellata,    115 

Proskauer,  case  of  Oxyuridm  in  nose,  696 
Prostomata,  230 

Protargol  in  balantidian  dysentery,  637 
Proteid  destruction  in  ancylostomiasis,  647 

—  metabolism  in  anaemia,  645 
Proteosoma,  spread  of  malaria  in  birds  by, 

158 
Protista  defined,  29 

—  spirochsetes  classed  among,  115 
Protomonadina,  52,   60 

—  classification,  60,  61 


882 


THE   ANIMAL    PARASITES    OF    MAN 


Protozoa,   25,   756 

—  alternation  of  generations  in,  27 

—  blood-inhabiting,  examination  of,  747 

—  characters,  25 

—  chromodial  apparatus  of,  26 

—  classification,   27 

—  clinical  and  therapeutical  notes  relating 

to,   617 

—  cytological    details,    method    of    examin- 

ing,  748 

—  definition  of,  25 

—  digestive  apparatus,  26 

—  ectoplasm   and  endoplasm   of,   25,  26 

—  encystment  of,  27 

—  examination,  methods  for,  745,  746 

—  food  of,  26 

—  genera   of,   precise    definition    sometimes 

impossible,  733 

—  hereditary  transmission  of  (footnote),  19 

—  intra   vitam    staining   of   fresh   prepara- 

tions, 746 

—  nucleus  of,  26,  27 

—  organellae,    29 

—  parasitic  in  blood,  culture  media  Ltr,  744 

—  propagation  of,  27 

—  sectioning  tissue  parasitized  by,  749 

—  or     bacteria,      question    whether    spiro- 

chsetes  to  be  classed  among,  115 
Protozoology,  notes  on  technique,  745-752 

—  recent  researches  in,  733 
Prowazek,  balantidian  dysentery,  637 

—  Chlamydozoa,   207 

—  Entamceha  biltschlii,  &c.,  34 
buccalis,  43 

—  Herpetomonas  muscce  domesticm,   102 

—  lamblial   diarrhoea,   625 

—  variety   of   Trichomonas   intestinalis   in- 

habiting oral  cavity,  56 

—  and  Aragao,  filtration  experiments  with 

chlamydozoal   granules,   209 
Prowazekia,  characters  of,  63 

—  asiatica,  65 

—  cruzi,  characters,  66 

—  javanensis,  characters,  66 

—  parva,    66 

—  urinaria,  63 

characters,  63 

flagellate  stage,  64 

in  cultures   associated  with  bacteria, 

65 
synonyms,  63,  64 

—  weinbergi,  characters,  66 
Prowazek's  bodies,  208 

Pruner,   Porocephalus   constrictus,   526,   527 
Pruritus  ani  due  to  escape  of  ascarides,  688 

set  up  by  migration  of  Oxyuris  ver- 

rnicularis,  695 

—  nematode     larvae     in     blood     associated 

with,  378 
Prussia,     oxen     infected    with     Cysticercus 
bovis  in,  341 

—  percentage   of   pigs   infected  with   cysti- 

cerci  in,  334 

—  proportion  of  trichinous  to  healthy  pigs 

in,  429,  430 
Pseudo-helminthes,  8 


Pseudomeningitis  due  to  Ascaridoe  infec- 
tion, 649,  650 

Pseudo-myxoedematous  form  of  Brazilian 
trypanosomiasis,  88 

Pseudonavicellae,  129,   1.30 

—  amoeboid  germs  in,  130 
Pseudoneuroptera,  characters,  531 
Pseudo-parasites,  6,  8 
Pseudophyllidea,   morphology,   308 
Pseudotaeniorhynchus,  576 

—  characters,  564 
Psorophora,  characters,  563,  571 

—  ovum  of,  557,  558 
Psoroptes,  characters,  517 
Psorospermia  of  Arthi'opoda ,   184 
Psorosperms    (Myxosporidia),    discovery    of, 

181 

—  egg-shaped,    former    name    for    Coccidia, 

135 

Psychodidce  (owl  midges),  moth-like  appear- 
ance of,  581 

PsychodinoR,  characters,  581 

Pterocephalus,  host  of,  135 

Pterygota,   classification,   531 

Pulex,  distinctive  characters,  545 

—  irritans  (human  flea),  bite  of,  effects,  714 
treatment,  714 

carrier  of  plague  bacillus,  543 

characters,  545 

larva  of,  546 

raaj  transmit  Trypanosoma  lewisi,  92 

—  pallipes,  548 

—  serraticeps  (dog  flea),  546 
Pulicidw  (true  fleas),  characters,  543 

—  classification  of  genera,  545 
Pulmonary  artery,  ascarides  in,  656 
Pumpkin  seeds  as  vermifuge,  673 
Pupipara  or  Eproboscidce,  blood-sucking,  611 
Purgatives  for  expulsion  of  ascarides,  693 

—  in  arrest  of  development  of  trichinosis, 

681 

Pustules  arising  from  clothes  louse  infec- 
tion, 711 

Putnam,   Oxyuris   in   appendix,   654 

Pycnosoma,  characters  of,  588 

—  and     Chrysomyia,     distinguishing     fea- 

tures, 588 

—  putorium,    spread    of   amoebic   dysentery 

by,  614 
Pyelitis  following  invasion  by  Eustrongylus 

gig  as,  682 
Pygiopsylla,  distinctive  characters,  545 
Pyorrhoea    alveolaris,    association    of    Ent- 

amoeba  buccalis  with,  43,  734 

of  species  of  Treponema  with,  128 

treatment,  620 

Pyretophorus,   characters,   561,   567 
Pyronin    producing    blepharoplastless     try- 

panosomes   (T.   brucei),   101 


Quincke  and  Eoos,  species  of  amoebae  named 

by,  31 
Quinine,      administration      as      preventive 

against  malaria,  636 


INDEX 


883 


Quinine,  administration  in  malaria,  635 

dosage,  635 

methods  of  administration,  635 

time   for,   635 

treatment   by,   635 

—  in  Indian  kala-azar,  626 

—  lotion,    irrigation    of    lower    bowel    with, 

in  gangrenous  dysentery,  619 


Rabbit,  development  of  hydatid  scolices  in, 
353 

—  host  of  Eimeria  stiedce,  145 

—  intestinal  coccidiosis  in,  145,  147 

- —  intestine  of,  section  infected  by  Eimeria 
stiedce,  145 

—  kidney    of,    use    in    cultivation    of    Tre- 

ponema pallidum,  126 

—  liver  of,  section  through  nodule  infected 

by  Eimeria  stiedce,  147 

—  Sarcosporidia  in,   187 

Eabies,  parasite  of,  cultivation,  210 
Eadiolaria,  characters  and  habitat,  28 
Radium  treatment  of  Oriental  sore,  628 
Railliet,  method  of  infection  with  Trichuris 

depressiuscula,  420 
Rainey's   corpuscles,   189 
Rain-water    barrels,    mosquitoes    depositing 

ova  in,  553,  557 
Ramstedt,   Oxyuris  infection   in   relation   to 

appendicitis,   653 
Ranken,  treatment  of  sleeping  sickness  with 

antimony,  623 
Basahus  bigiittatus,  bite  of,  540 

—  —  geographical  distribution,  540 

—  —  synonyms,  540 

Rat  attacked  by  Dermatophilus  (Sarco- 
psylla)  penetrans,  613 

—  blood    of,    transference    of    Trypanosoma 

brucei  from,  to  blood  of  snake,   102 

—  blood  parasite,  see  Trypanosoma  lewisi 

—  gut    and    caecum    of.    Trichomonas    from, 

735 

—  infection    with    Trichinella,    method    of, 

427 

with  Trichinella  spiralis  in  slaughter- 
houses and  knackers'  yards,  427 

with  Trypanosoma  lewisi,  mode  of,  92, 

93 

—  muscles      of,      invaded      by      Trichinella 

spiralis,   425 

—  normal  host  of  Trichinella  spiralis,  427 

—  pneumocysts  in,  90 

—  sewer   and   black,   hosts   of   Hymenolepis 

diminuta,   326 

—  flea    (Ceratophyllus    fasciatus),    cysticer- 

coid  of  Hymenolepis  diminuta  found  in, 

327,  328 

host  of  rat  trypanosome,  88,  90,  543 

larval  stages  of  Hymenolepis  murina 

occurring  in,  17 
see   also  Ceratophyllus  fasciatus 

—  Trypanosoma  lewisi  in,  88 


Rectum,    administration    of    quinine    by,    in 
malaria,  635 

—  bilharziasis  of,  642 
treatment,  644 

—  irrigation   of,   in   intestinal   myiasis,   728 

—  means    of   access    of   Schistosoma   hasma- 

tobium  to,  272 

—  pathological  changes  in,  due  to  Schisto- 

soma hcematobium,  274,  275 

—  plexus    formed    in,    by    superior    haemor- 

rhoidal  veins,  272 
Redi,  origin  of  flesh  maggots,  10 
Redise  of  trematodes,  225,  226,  227,  228 
Reduviidce,  bites  of,  537 

—  characters  of,  537 

—  geographical  distribution,  537 
Reduvius    personatus,    bite    of,     sometimes 

fatal,  539 

geographical  distribution,  539 

Red-water  fever,  European,  in  cattle,  cause 

of,  177 
Reighardia,  523 
Relapsing  fever,  120,  629 

African,  cause  of,  116,  630 

incubation  period,  630 

prophylaxis,   631 

symptoms,  630,  631 

—  treatment,  631 

American,    630 

Asiatic,  mortality  from,  631 

prophylactic  measures,  631 

symptoms,   631 

treatment,  631 

complications,  630 

East  African,  cause  of,  122 

European,   agent  of,    122,   629 

incubation    period,    630 

prophylaxis,    630 

symptoms,   630 

treatment,    630 

Indian,  cause  of,  122 

North  African,  prophylactic  measures, 

631 

symptoms,  631 

—  treatment,  631 

and  Egyptian,  cause  of,  122,  631 

—  —  prophylactic  measures,   630 

spirochaetes    causing,    115,     120,    122, 

508 

transmission  by  lice,  120,  630 

by  ticks,  117,  630,  631 

treatment,   630 

Remak,  investigations  of  Coccidia,   135 
Reptiles,  haemogregarines  in,  153,  154 

—  Sarcosporidia  in,  187 

Resorcin  ointment,   application   in   creeping 

disease,  732 
Respiration,   organs   of,    in   Insecta,    530 
Retinal       haemorrhages       in       ancylostome 

anaemia,  646 
Reyher,  bothriocephalus  anaemia,  644,  645 
Rhabdites     in     gastric    fluid    obtained    by 

lavage,  378 

—  mellio,  presence  in  vagina,  377 

—  niellyi,   378 

mode  of  infection   in  man,  378 


884 


THE   ANIMAL    PARASITES    OF    MAN 


Bhahditis    pellio,    induction    of    facultative 

parasitism,  377 

morphology,  377 

synonyms,   377 

Ehabdonema,    alternation   of   parasitic   and 

free-living  generations,  20 

—  life-history  of,   19 

—  propagation      of,      parasitic     generation 

during  free  life,  18 
Rheins,  case  of   Oxyuridce  in  nose,  696 
Rhinosporidium,  195 

—  hosts  of,   197 

—  in  conjunctival  polypus,   197 

—  in  external  auditory  meatus,  196 

—  in  horses,   197 

—  in   nasal   polypus,    195 

—  in  papilloma  of  penis,  197 

—  kinealyi  (or  seeberi),  195,  197 

causal  agent  of  a  nasal  polypus,  195, 

196 

cysts  of,   196 

geographical    distribution,    195,    196 

pansporoblasts  of,  196 

trophozoites  of,  196 

tumours  produced  by,  197 

Rhipicentor,  characters  of,  497 
BhipicephalcB ,  characters  of,  496,  497 
Rhipicephalus,  characters  of,  497 

—  species  of,  transmitting  Theileria  parva, 

179 

—  annulatus,    carrier    of    Texas     fever    in 

cattle,  494 
■ moulting  of,  496 

—  appendiculatus    and    B.    simus,    carriers 

of  Rhodesian  fever  in  cattle,  494 

—  bursa,     trasmitting     agent     of     Babesia 

bovis,  177 

—  sanguineus,     geographical     distribution, 

505 

hosts  of,  505 

synonyms,  505 

—  —  transmission      of      leucocytogregarine 

from  dog  to  dog  by,   155 
transmitting  agent  of  Babesia  canis, 

177 
Bhizoglyphii,   characters   and  habitat,  514 
Bhizoglyphus  parasiticus ,  characters,  514,  515 

skin  disease  produced  by,  514 

Rhizopods,  flagella  occurring  among,  52 

—  possible    association    with    poliomyelitis 

acuta,  47 
Rhodesian   fever  in  cattle,  carriers  of,  494 
Bhodinus  prolixus,  bite  of,  541 

geographical  distribution,  542 

Bhyncobdellidce,  482 
Rhyncobothrium,  scolices  of,  305 
Bhyncota,   see  Hemiptera 
Bicinidm  classed  among  mutualists,  6 
Riley,  see  Walsh  and  Biley 
Rinderpest  and  coccidiosis,  741 

—  method  of  transmission,  742 

River  fever  set  up  by  kedani  mite  in  Japan, 

487 
Rivolta,  experimental  infection  with  Cocci- 

dia,  136 

—  on  Sarcocystis   lindemanni,   193 


Robertson,  Miss,  development  of  Trypano- 
soma gambiense,  74,  75 

in   Glossina  palpalis,  74,  75 

forms  of  Trypanosoma  gambiense,  73, 

737 

Rocky  Mountain  spotted  fdver,  carrier  of, 
496 

tick  fever,  carrier  of,  503 

mortality,   504 

Rodenwaldt,  distribution  of  larvae  of  Filaria 
immitis  in  body,  393 

—  periodicity  of  larvae  of  Filaria  bancrofti 

in  peripheral  blood,  393 
Rogers,    Sir    L.,    agent    transmitting    kala- 

azar,  713 
cultivation     of    Leishmania    dono- 

vani,    105,    106 
places  Leishmann-Donovan  body  in 

genus  Herpetomonas,  107 
transmission  of  surra  by  Chrysops, 

601 
treatment    of     amoebic     dysentery, 

618 

treatment  of  Indian  kala-azar,  626 

of  pyorrhoea  alveolaris,  620 

Rokitansky,     perforation     of     intestine     by 

Ascaris,  655 
Romani,  agglutinating  haemolytic  action  of 

serum  of  ancylostome  patients,  648 
Romanowsky  stain,   749 

slightly  modified,  formula  of,  750 

underlying  principle  of,  750 

Roos,  presence  of  cercomonads  in  gan- 
grenous lung,  62 

—  and     Harris,     penetration    of    intestinal 

blood-vessels  by  amoebae,  36 
Rosenquist,  proteid  metabolism  in  anaemia, 

645 
Ross,  E.   H.,  Treponema  pallidum,  124 
Ross,    Sir   Ronald,    campaign    against   mos- 
quitoes in  prevention  of  malaria,  636 

development    of    malarial    parasite 

in  mosquito  traced  in  Plasmodium  relic- 
turn  by,  171 

discovery       of       transmission       of 

malarial    parasites   by   mosquito,    158 

"  Prevention  of  Malaria,"  617,  633 

relapses  in  malarial  fever,  161,  162 

trichomonads  and  cercomonads,  56 

and  Thomson,  D.,  cyclical  varia- 
tion of  trypanosomes  in  blood,  78 

—  method     of     determining 

number  of  trypanosomes  in  blood,  747, 
748 
Rossia,  characters,  568 
Rossi^lla,  morphology,  174 

—  rossi,  174 
Rostellum  of  Cestoda,  289 

Rothschild,  classification  of  genera  of  Puli- 

cidcp,  545 
Roubaud,  cause  of  myiasis  in  French  West 

Africa,  614 

—  life-histo-v      of      Cordylobia      nnthropo- 

vhaga,  614 

—  Pycnosoma  putorium,   614 


INDEX 


885 


Eovelli,      larval      stage      of      Hymenolepis 

diminuta,   327 
Row,    experimental   production   of   Oriental 

sore,    109 
—  treatment  of  Oriental   sore,   628 
Rudolphi,  origin  of  helminthes,  12 
E-uffer,  lesions  produced  by  Oxyuris  vermi- 

cularis,  695 
Buncliiomyia,  characters,  565 


S. 


Sabadill  vinegar,  lotions  of,  in  head  louse 

infection,   710 
Sabatier  on  change  of  hosts,  21 
Sabethes,  characters,  565 
Sabethoides,  characters,  565 
Sachs,  treatment  of  scabies,  707 
Sack,  treatment  of  scabies,  707 
Sahara,   Central,   human   myiasis  occurring 

in  mountains  of,  598 
St.   Artault,   Trichomonas  pulmonalis,  56 
Saline  solution,  physiological,  lavages  of,  in 

myiasis,  719 
Salol  as  tapeworm  drug,  674 
Salt  water,  mosquito  larvae  living  in,  557 
Salvarsan    in   Asiatic   relapsing   fever,   631 

—  in  North  African  relapsing  fever,  631 

—  in  Oriental  sore,  628 

—  in   relapsing  fever,   630 

—  in  tropical  syphilis,  632 

—  in    trypanosomiasis,    623 

—  in  yaws,  632 
dosage,  632 

Salzmann,   mode    of   infection    in    intestinal 

myiasis,  727 
Sambon,  L.  W.,  Linguatula  serrata,  527 
personal   experiments   with  regard  to 

malarial   infection,   158 
Samelsohn,  retinal  haemorrhages  in  ancylo- 

stome   anaemia,  646 
Sandal    oil    in    chyluria   from    Filaria    han- 

crofti  infection,  677 
Sand  flea,   see  Dermatophilus   (Sarcopsylla) 

penetrans 

—  flies,  577 

and    fever    due    to    them    in    North 

China,    613 

biting  in   Hampshire,   579 

Sandflies,  haunts  of,  613 

see  also  Siynulium 

Sandler,  trichocephalus  anaemia,  651 
Sandwith,  F.  M.,  toxic  symptoms  following 

thymol  administration,  686 

treatment  of  bilharziasis,  643 

Santonin  in  bilharziasis,  643 

—  in  expulsion  of  ascarides,  692 
^  of  OxifuridcB,  697 

—  in    intestinal    myiasis,    728 

Sapo  viridis  and  tar,  application  in   creep- 
ing disease,  732 
Sarcocystin,  isolation  of,  191 
Sarcocystis  hertrami,  193 

—  blanchardi,   193 
from   ox,    190 


Sarcocystis  colii,   193 
spore,   193 

—  hueti,  193 

—  lindemanni,  193 

—  miescheriana,  188,  193 
from   pig,   190 

—  muris,   193 

deadly  to  host,  191 

experimental  infection  with,   191,   192 

gymnospores  of,   191 

spore  of,  site  of  sarcocystin,  192 

—  of  muscles,  191 

—  pansporoblasts    of,    189 

—  recognition    from    other    foreign    bodies, 

188 

—  spores  of,   189 

—  tenella,  193 
from  sheep,   190 

—  —  spores  of,  191 

toxin   isolated   fronl,   191 

Sarcodina,  27,  29 

—  characters  and  habitat,  27 

Sarcoid    globules    in    miracidium    of    Schis- 

tosoma  hcBmatobium,  276 
Sarcophaga   carnosa    (flesh  fly),   characters, 

589 

larvae  of,  589 

regions    of    human    body    invaded 

by,  589 
viviparous,   589 

—  chrysostoma,  590 

—  hcematodes,  589 

—  hoBniorrhoidalis ,  589 

—  magnifica,  geographical  distribution ,  589 
larvae  of,  regions  of  human  body  in- 
vaded by,  589 

references  to,  589 

—  plinthopyga,  probably  concerned  in  dis- 

semination of  yaws,  590 

—  ruficomis,  589 

—  wohlfahrti,  larvae  of,  method  of  destroy- 

ing,  723 

unusual  situations  of,  723 

nasal  myiasis  from,  722,  723 

Sarcopsyllidoe,   characters,   543 
Sarcoptes,   characters,   517 

—  species      transmissible      from      domestic 

animals  to  man,  520 

—  auchenii,  520 

—  canis,  520 

—  dromedarii ,  520 

—  equi,  520 

—  leonis,  520 

—  minor,  hosts  of,  520 

—  ovis,  520 

—  scahiei,   characters,  518 

infection  by,  disease  produced  by,  704 

see  also  Scabies 

synonyms,  518 

crnstosce,  519 

var.  hominis,  519 

development  of,  519 

excavation  of  tunnels  in  human 

epidermis   by,   517,   519 
■  transmission     of,     natural     and 

artificial,  519 


886 


THE   ANIMAL    PARASITES    OF    MAN 


Sarcoptes  suis,  520 

—  vulpis,  520 

Sarcoptidce   (itch  mites),   characters,  516 

—  development,  stages  in  male  and  female, 

517 

—  hosts  of,  516 

—  rate  of  breeding,  517 
Sarcoptince,  517,  518 
Sarcosporidia,  129,  187,  193 

—  chambers   of,   189 

—  characters  and  habitat,  28,    188 

—  experimental   transmission,   191,    192 

—  fatal  to  sheep,   188 

—  hosts  of,   187 

—  in  man,   193 

—  morphology,  188 

—  muscles  affected,   188 
Sarcosporidiosis,      possible      percentage      of 

animals  affected  by,   191 
Scabies,  704 

—  diagnosis,  705 

from  occupational  eczema,  706 

—  mite  tracks   of,   705 

—  prognosis,   706 

—  symptoms  of,  705 

—  treatment,  706,   707 

—  norvegica  (Norway  itch),  520,  705 
Scabiophilia,  706 

Scarlet  fever,  cell  inclusions  in,  208 
Schaudinn,       classification       of      intestinal 
amoebae,    31 

—  cytological    changes    during    encystment 

process  of  Entamoeba  coli,  33 

—  infection    by   trichomonads,    56 

—  intensity    of    infection    with    Entamoeba 

coli,  33 

—  on  Leydenia  genimipara,  49 

—  on   Paranioeba   hominis,   44 

—  penetration    of   red    blood    corpuscles    by 

sporozoites  of  tertian  parasite,  159 

—  relapses  in   malarial   fever,   161 

—  researches  on  Coccidia,  137,  138,  139 
on   Entamoeba   histolytica,   34,   37 

—  Treponema  pallidum,  114 
Schaudinn's  fluid,  748 
Scheube,  lung-fluke  disease,  639 
Schewiakoff,   movements   of  gregarines,   131 
Schiller,   Ascaris   and   Oxyuris   infection   in 

relation  to  appendicitis,  653 

—  effects  of  trichocephalus  infection,  651 
Schistocephalus,  pleroceroid  of,  300 
Schistosoma,   morphology,   269 

—  synonyms,  269 

—  cercariae,  753 

—  hmmatobium,  distribution   in  body,  272 

endoiihlebitis  set  up  by,  274,  275 

female,    diameter,    273,   274 

morphology,  271 

genitalia,    276 

geographical   distribution,  276 

in   caval   system,   274 

in   gall-bladder,  274 

in  hsemorrhoidal  veins,  273 

in  lungs,  274 


Schistosoma     hcematobium     in     vesico-pros- 

tatic  plexus,   273,   274 

infection   hj,   641 

see   also   Bilharziasis 

male,  anterior  end,  diagram  showing 

organs,  271 
carrying   female   in   canalis   gynae- 

cophorus,   270 

diameter,  273 

morphology,    270 

and    female    in    copula,    transverse 

section  through,  271 
means  of  access  to  descending  colon, 

rectum,  anal  canal,  bladder  and  caval 

system,  272 
miracidium    of,    sarcoid    globules    in,. 

276 
most    easily    found    post    mortem    in 

portal  vein  and  liver,  273 

ova  in  utero,  diameter,  273 

lateral  spined,  273 

origin  of,  273 

ovum  of,  277 

pathological    changes    in    rectum    and 

bladder  due  to,  274,  275 
synonyms,   270 

—  japonicum,  277 

anterior    end    with    testes,    posterior 

end  with  point  of  union  of  caeca,  278 

—  —  female,  morphology,  278 

from   dog,  280 

egg  from  faeces,  280 

uterine  egg,  280 

habitat,  280 

liver  showing  eggs  in  the  intra-  and 

interlobular  connective  tissue,  282 

male,  morphology,  277 

— ^  and  female  in  copula,  279 

—  —  mode  of  infection   by,   279 
ova  of,  278 

from        human        liver,        showing 

"spines"  and  "hoods"  at  opposite 
pole,  279 

sites  in  which  found  in   body,  282" 

—  mansoni,  754 
Schistosomidce,    269,    753 

[    —  morphology,   233 

I    Schizocystis,   135 

j    Schizogony  absent  in  Eugregarinea,  134 

i    —  in  Coccidiidea,  138 

—  in  Leucociitozoa,   153 

—  of   malarial    parasites,    161,    172 
Schizogregarinea,   135 
Schizotrypanum  cruzi,  83 

Schleip,  blood  examination   in  diagnosis  of 
I  trichinosis,  681 

j    Schlesinger,   intestinal   myiasis,   727 
Schliiter,       haemorrhagic       enteritis       from 

Strongyloides   infection,  674 
Schmidt,   larvae   in   nose   in    enormous   num- 
bers, 716 

—  Trichomonas  pulmonalis ,  56 
Schneider,  A.,  on  Coccidia,  137 
on    Eimeria,    142 

on   gregarines,    130 


INDEX 


887 


Schuberg,   copulation   in   Coccidia,   137 

—  immunity      of      Ornithodorus      moubata 

against  infection   with  Spirochceta  dut- 

toni,   119 
Schiiffner,       peculiar       fever       resembling 

typhoid,  613 
Schiiffner's  dots,   165,   166,   171 
Schultz,  on  Coccidia  in  cattle,  741 
Schupfer,   typhoid   lumbricosis,  650 
Schwankhaus,  Ascaris  infection  in  relation 

to  appendicitis,  653 
Schweriner     itch     following     infection     by 

ancylostomes,  684 
Schwetz,      life-history      of      Auchmeromyia 

luteola,  614 
Scolex   of   tapeworms,   300,    303,   304 

morphology,  304 

Scolopendra      in      maxillary      and      frontal 

sinuses,  721 
Screw  worm,  Indian,  see  Pycnosoma 
fly,     see     Chrysomyia     (Compsomyia) 

macellaria 
Scutomyia,  characters,  563 
Seal,  host  of  Dibothriocephalus  latus,  315 
Sebirol  as  vermicide,  672 
Seeber,   Ehinosporidium  described  by,   197 
Sehrt,    abscess    of    omentum    with    Ascaris 

ova   in   pus,   657 
Seidelin,   association   of  Paraplasma  bodies 

with  yellow  fever,   180 
Seifert,       blood-stained       diarrhoea       from 

Strongyloides  stercoralis  infection,  674 
Selenidium,  135 

Sellards,  see   Walker  and  Sellards 
Senevet,    herpetomonad    flagellate    in    cul- 
tures of  blood  and  organs  of  geckos,  739 
Senna,    syrup    of,    in    expulsion    of    ancylo- 
stomes, 686 
Sense,     organs     of,     lacking     in     parasitic 

nematodes,  366 
Sepsidce,  characters,  583 

—  larvae  (maggots)  of,  583 
Septicaemia      terminating      nasal      myiasis 

fatally,  718 
Sergent,    transmission    of    relapsing    fever, 

120,  121 
Sergent,  Ed.  and  Et.,  herpetomonad  flagel- 
late in  cultures  of  blood  and  organs  of 

gecko,  739 

''thymni,"  725 

Sergent,    E.    and   L.,    deposition    of   ova   of 

Oestrus  ovis,  598 
transmission    of    trypanosomes    by 

species  of  Tabanus,  601 
Sergent    and    Gillot,    treatment    of    North 

African  relapsing  fever,  631 
,  Sergentella  hominis,  210 
Serous     fluid,     bodies     resembling     amoebae 

found  in,  46 
Serum  diagnosis  of  echinococcus,  359 

complement   deviation,   359 

precipitin  reaction,  359 

—  human,    action    on    Trypanosoma    rhode- 

siense,   80 

—  immune,   action   on    Trypanosoma   rhode- 

siense,   80 


Setaria,  habitat,  407 

—  morphology,  407 

—  equina,   hosts   and  habitat  of,   408 
morphology,    408 

synonyms,    408 

—  (Filaria)  hcemorrhagica,  408 

—  labiata  papillosa,  408 

Sexual  organs,  irritative  effects  on,  set  up 
by  migrations  of  Oxyuris  vermicularis, 
695 

of  Echinorhynchus,  476 

of  Hirudinea,   481 

of   Insecta,   530 

of  nematodes,  367,  368,  369 

Sheep,  baleri  in,  causal  agent,  95 

—  "  carceag "  in,  cause  of,  177 

—  Cysticercus   cellulosoe  in,  337 

—  echinococci   in,   346 

—  heart-water   fever   in,   carrier  of,   493 

—  how  infected  by  Fasciola  hepatica,  226 

—  liver-fluke    disease    in,    death    from    apo- 

plexy in  first  period,  240 

period  of  anaemia,  240 

of    immigration,    240 

— of  migration  of  flukes,  241 

of  wasting,  240 

ravages   caused  by,   238 

—  organs  infected  with  echinococcus,   per- 

centage  of   frequency,   347 

—  Sarcosporidia  fatal   to,   188 

—  section  of  Sarcocystis  tenella  from,  190 
Sheep-ked,  see  Melophagus  ovinus 

Shell   gland   secretion   in  trematodes,   223 
Shiga,  discovery  of  dysentery  bacillus,  31 

—  species     of     amoebae     distinguished     by, 

31 
Shipley,  A.  E.,  prophylaxis  against  clothes 

lice,  616 
Sick,  cases  of  ascarides  in  bile-ducts,  688 
Siebert,  application  of  epicarin  in  scabies, 

707 
Siebold,  v.,  development  of  Taeniae,  14 

—  explanation  of  bladder  worms,   14 

—  feeding  experiments  with  Tcenia  echino- 

coccus, 356 

—  investigations  of  Gregarinida,  129 

—  observation  of  Pseudonavicellae,  129 

—  psorosperms,  181 

—  views  as  to  development  of  Helminthes, 

13 
Siedlecki,  researches  on  Coccidia,  137 
Siegel,  Cytorhyctes,  208 

—  Cytorhyctes   luis,   124,   208 

Silcock,  case  of  human  hepatic  coccidiosis, 

148 
Silkworm   disease,    "  gelbsucht,"    207 

Nosema  cause  of,   184 

Silvanus      surinamensis ,      characters      and 

habitat,  543 
Silver  tick,  see  Amblyomm,a  cayennense 
Simond,  researches  on  Coccidia,  137 
Simulidce,  577 
Simulium,  bite  of  species  of,  578,  579 

—  characters,    577 

—  larvae  of,  578 

—  life-cycle  of,  578 


888 


THE   ANIMAL    PARASITES    OF    MAN 


Simulium,   wing  of,  579 

—  buissoni,  possible  connection  with  spread 

of  leprosy,  579 

—  columoascnensis ,    geograjjhical    distribu- 

tion, 578 

—  damnosum,  geographical  distribution,  578 

—  griseicoLLis,     geographical     distribution, 

579 

—  latipes,   579 

—  meridionale,  possible   carrier  of  chicken 
cholera,   579 

—  occidentalis,  579 

—  wellmanni,  579 

Sinton,    culture    of    trypanosome    forms    of 

T.  gamhiense,  76 
of  T.  rhodesiense,  83 

—  Prowazekia  urinaria,  64,  65 
Siphunciilata,  532 

—  see  also  Pediculidoi 

Skin  affections  caused  by  cereal  mites,  489 

due    to   Dermanyssiis    hirundinis ,    492 

set   up  by  Leptus   autiimnalis,   reme- 
dies against,  702 
by    Trombidium    tlalsahuate,    486 

—  disease   caused  by  larvae  of  Dermatobia 

noxialis,   725 

due  to  young  nematodes  in  dogs,  378 

produced  by  Bhizoglyphus  parasi- 
ticus, 514 

—  diseases  set  up  by  penetration  of  larvae 

of  Ancylostoma  duodenale,  455 
various  names  for,  455 

—  filaria  infection  of,  378 

—  infection     by     Ancylostoma     duodenale 

through,  683 

by  larvae  of  Ancylostoma  duodenale, 

454,  455 

—  lesions  due  to  Sparganum  mansoni,  318 

—  mole,  599 

—  parasites    of    dogs     and    cats    infecting 

them   with  Dipylidium   caninum,   323 

—  surface  of,  larvae  on,  721,  722 
Skusea,  characters,   563 
Slaughter-houses,     infection    of    rats    with 

Trichinella  in,  427 

Sleeping   sickness,   68,   69,   72,   76,   620 

association  of  trypanosomes   with,   68 

cerebral   stage,  621 

due   to    Trypanosoma    gambiense,    68, 

72,  620 

rhodesiense,  69,  76,  620 

symptoms,  622 

febrile  or  glandular  stage,  621 

incubation  period,  621 

investigation   of,  68 

parasites  producing,  72,  76,  605 

pathology  of,  621 

preventive  measures,  623 

Rhodesian,  daily  number  of  trypano- 
somes in  blood  from  case  of,  79 

transmission  of,  68,  605,  607,  608 

experimental    (with   apes),   68 

treatment    by    arsenic    and    arsenical 

preparations,  622,  623 

by    atoxyl,    622 


Sleeping  sickness,  treatment  by  tartar 
emetic,  622 

must      be      commenced      in      early 

stages  to  be  effective,  622 

Sleeping  Sickness  Bureau  Bulletin,  founda- 
tion of,  69 

Sloth,  blood  of,  inhabited  by  Endotry- 
parium  schaudinni,  99 

Smith,  Theobald,  experimental  infection  of 
mice  with  Sarcocystis  maris,   191 

—  and  Barrett,  Endamoeba,  734 
Endamoeba  gingivaiis,  733 

treatment  of  oral  endamoebiasis,  620 

Smith  and  Kilborne,   174,   176,  177 

—  and  Weidman,  Entamoeba  mortinatalium , 

45 
Smithia,   morphology,    174 

—  microti,   174 

—  talpcB,    174 

Snake,    blood   of,   transference   of   Trypano- 
soma brucei  to,  from  blood  of  rat,   102 
Soamin  in  sleeping  sickness,   623 
Soda,  bicarbonate,  with  iodoform  in  expul- 
sion  of   ascarides,   694 

—  salicylate  of,  lavages  of,  in  nasal  mvia- 

sis,  719 
Soldiers,   Pediculus   vestimenti  pest   among, 
during  campaigns,   533 

—  prophylaxis   against  clothes  lice  among, 

616 

Solium,  derivation  of  specific  term  (foot- 
note), 331 

Souma  in  bovines  and  equines,  causal  agent, 
100 

Space  parasites,  20 

Spaniopsis   tabaniformis ,   614 

Sparganum,  317 

—  mansoni,    317 
cephalic  end,  318 

diagnostic    signs    of   j^resence,    659 

discovery  of,  317 

geographical  distribution,  659 

habitat  in  body  of  man,  659 

migration  in  body,  318 

plerocercoid   of,    318 

skin  lesions  due  to,  318 

;  symptoms  set  up  by  invasion,  059 

synonyms,  317 

transverse  section  of,  318 

—  proliferum,    318 

acne-like  condition  set  up  by,  318 

geographical  distribution,  320 

mode  of  infection,   320 

morphology,  319 

synonyms,   318 

Spengel,  Filaria  (?)  kilimaroe,  407 

Spermatozoa  of  Trematoda,  no  essential 
difference  in  structure  from  those  of 
other   animals,   222 

Sphcerularia,  nematodes  hatched  from  eggs 
of,  5 

Spiders,  see  Arachnoidea ,  483 

Spinal  ganglia  of  rabid  monkeys,  cultiva- 
tion, 210 

Spinning  mites,  see  Tetranychidcs 

Spirochaeta,  115 


INDEX 


889 


Spirochoeta    aboriginalis,     association     with 
granuloma  inguinale,    122 

—  acuminata,    122,    128 

—  anodontce,    114 

—  anserinu,  119,  122 

—  balbiunii,   114 

—  berberu,    agent    of    North    African    and 

Egyptian  relapsing  fever,   122 

—  bvonchialis,    122,    632,    739 
mode   of   infection,   740 

morjihology  and  life-history,  739,  740 

—  buccalis,  122 
morphology,   741 

—  cavtcri,  agent  of  Indian  relapsing  fever, 

122 

—  dentium,    122,    128 
morphology,   741 

—  duttoni,   116 

agent  transmitting,  116 

cause  of  African  relapsing  fever,  116, 

630 

cultivation  of,   123 

geographical   distribution,   116,    119 

infection   by,  experimental,  117 

summary    of    methods    and    re- 
sults,   118,    119 

—  —  — ■  immunity    of    Ornithodorus    mou- 

bata  against,  119 
transmission  of,  739 

—  eurygyraia,   122 

—  galiinarum,   119,  122 

agent  of  transmission,  119 

appearance     in     hsemocoelic     fluid     of 

Argas  yersicus,  119     • 

cultivation  of,   123 

fatal  to  fowls,  119 

—  gigantea,   114 

—  granulosa,   116 

—  hachaizce  in   cholera  motions,    122 

—  laverani,  small  size  of,   122 

—  marchouxi,    see    Spirochceta   galiinarum 

—  muris,   122 

—  novyi,  agent  of  North  American  relaps- 

ing fever,  122 
cultivation  of,   123 

—  obermeieri,  see  Spirochceta  recurrentis 

—  obtusa,   122,    128 

—  ovina,   122 

—  phagedenis,   122 

—  2}licatilis,  114 

—  recurrentis ,  120 

agents  of  transmission,  120 

cause    of    European    relapsing    fever, 

120,  122 

■ cultivation   of,   123 

incubation  period,  630 

morphology,  120 

—  ref ring  ens,    122,    128 

association  with  Treponema  jyalUdum., 

122 

—  rossii,   agent   of   East   African   relapsing 

fever,   122 
cultivation  of,  123 

—  schaudinni,  agent  of  ulcus  tropicum,  122 

—  stenogyrata,  122 

—  theileri,  122 


Spirochceta  vincenti,   122 
Spirochcetacea,   115 
Spirochsetes,  114 

—  blood  inhabiting,    116 

—  classed  among  Frotista,  29,  115 

—  cultivation   01,    123 

presence  of  oxygen  necessary  for,   123 

—  granule   phase   of,    120 

—  hosts  of,   114 

—  in   alimentary  tract,   741 

—  in    human   mouth,    122,   740 

—  in  voinited  matter,   122 

—  mode  of  division,  115 

—  molluscan,    breaking    up    into    granules, 

119 

—  morphology     and     morphological     varia- 

tion, 114,  115 

—  of   human   mouth,   recent   Avork   on,    740, 

741 

—  of  relapsing  fever,  periodic  increase  and 

decrease  in  blood,   115 

—  reaction  to  drugs,  115 

—  systematic  position,   115 
Spirochcetoidea,   115 
Spirochsetoses,   629 

—  bronchial,   diseases  for   which  mistaken, 

632 
treatment,   633 

—  relapsing  fever,  629 

—  syphilis,   632 

—  yaws,  632 
Spiroschaudinnia,   115 

Spleen,  development  of  crescents  of  tertian 
malignant  parasite  in,  169 

—  enlargement  due  to  ova  of  Schistosoma 

japonicum,    282 
in  malaria,  634 

—  pigmentation      of,      following      malaria 

(footnote),  165 
Splenic     blood,      citrated,     cultivation     of 
Leishmania  donovani  in,  106 

—  vein,  tributary  of  portal  vein,  272 
Splenomegaly,    association    of    Histoplasma 

capsulatum.  with,  112 
of  Toxoplasma  pyrogenes  with,  113 

—  infantile   (kala-azar),  109,  627 
Spontaneous  generation,  theory  of,   10 

early  opposition  to,   10 

Sporoblasts  of  Coccidiidea ,  141 

—  of  malarial  parasites,   163 

—  of  Myxosporidia,    183 
Sporocyst,  germ  balls  of,   227 

—  of  Coccidiidea,   141 

—  of  gregarines,  134 

—  of  trematodes,  225,  227 
Sporogony,  144,  186 
Sporozoa,  28,   128 

—  characters   and  habitat,  28 

—  classification,    129 

—  hosts  of,   129 

—  relation   to   Protozoa,   19 

I    Sporozoites  of  Coccidiidea,  138,  139,  140 

i    —  of  gregarines,  132,  133 

'    —  of  malarial  parasites,  159 

j    Stained  material,  examination  of,  747 

'    Staining,   749 


890 


THE   ANIMAL   PARASITES    OF    MAN 


Stallion's    disease    (dourine),    trypanosomes 

in  blood  of  horses  with,  68 
Stannus,    species   of   EnyaLiopsis   producing 

ulcers,  542 

—  and  Yorke,  observation  of  Trypanosoma 

rhodesiense  in  animals  inoculated  from 
case  of  sleeping  sickness,  78 

Staphylocystis,   304 

Staubli,  blood  examination  in  diagnosis  of 
trichinosis,  681 

Steel  and  Evans,  experimental  transmission 
of  Trypanosoma  evansi,  67 

Steenstrup,  discovery  of  method  of  multi- 
plication  of  Heiminthes,   13 

Stegomyia,  breeding  of,  prevention,  636 

—  characters,  563,  571 

—  ovum  of,   557,   558 

—  transmission  of  yellow  fever  by,  555 

—  albipes,  characters,  572 

—  albocephala,  characters,  573 

—  albolateralis ,  characters,  573 

—  albomarginata,  characters,  573 

—  amesii,  characters,  573 

—  argenteomaculata,  characters,  572 

—  argenteopunctata,   characters,   573 

—  assamensis,  characters,  573 

—  auriostriata,  characters,  573 

—  crassipes,  characters,  573 

—  dubia,  characters,  573 

—  fasciata,  biting  hours  of,  574 
breeding  of,  574 

carrier  of  yellow  fever,  574 

characters,  572,  574 

development  of  Plasmodium  relictum 

in,  171 

—  —  distinguishing    characters   of   S.    scu- 

tellaris  from,  575 

domesticated  species,  574 

food  of,  574 

geographical   distribution,    574 

larvae  of,  habitat,  574 

ova  of,  574 

possible   host  of  Leishmania   tropica, 

108 

source   of   danger  to   Panama   Canal, 

574 

supposed  intermediate  host  of  para- 
site of  Bagdad  sore,  575 

transportation  of,  574 

—  gelebinensis,  characters,  572 

—  grantii,  characters,  573 

—  lilii,  characters,  572 

—  mediopunctata,  characters,  573 

—  minuta,  characters,  573 

—  minutissima,  characters,  572 

—  nigeria,  characters,   572 

—  poweri,  characters,  572 

—  pseudonigeria,  characters,  572 

—  pseudonivea,  characters,  573 

—  pseudoscutellaris ,  394,  575 
characters,  572 

intermediate  host  of  filaria  in  Fiji,  575 

—  punctolateralis ,   characters,   573 

—  scutellaris ,    characters,    572 

distinguishing  character  from  S.  fas- 
ciata, 575 


Stegomyia  simpsoni,  characters,  572 

—  terreus,   characters,   573 

—  tripunctata,   characters,  573 

—  W-alba,  characters,  572 

—  wellmannii,  characters,  572 

Stein,  interrelation  of  pseudonavicellae  and 

gregarines,   129 
Stein,  v.,  classification  of  Infusoria,  199 

—  discovery     of     meal     worm     in     bladder 

worm,   303 

Steinhaus,  intestinal  stenosis  following  in- 
fection by  Tcenia  solium,  662 

Stempell,  on  Nosema  bombycis,  184 

Stephens,  J.  W.  W.,  appendix  on  Trema- 
toda  and  Nematoda,  753 

Nemathelminthes,   360 

Plasmodium  tenue,  170 

Platyhelminthes    or   flat   worms,    211, 

638 

and  Christophers,  Maurer's  dots,  168 

relapses    and    latent    infection    of 

malaria,   158 

and  Fantham,  length  of  Trypanosoma 

gambiense,  73 

Trypanosoma   rhodesiense,   69,   76 

Stern,  symptoms  of  cysticercus  in  fourth 
ventricle,  665 

Stethomyia,  characters,  561,  567 

Stiles,  C.  W.,  infection  with  Lamblia  in- 
testinalis,  60 

prophylaxis  against  flagellate  diar- 
rhoea,  625 

Stillborn  child,  problematical  "  monocystid 
gregarines  "  from  lung  tissue  of,  150 

Stitt,  alkaloid  of  quinine  in  malaria,  635 

—  paroxysms    of    malignant    tertian    fever, 

634 
Stock,  bilharziasis,  641 

—  treatment  of  bilharziasis,  644 
Stokvis,     Balantidium     coli     occurring     in 

lung,  202 
Stomach,    cancer    of,    Lamblia    intestinalis 
in,  59,  60 

—  fluid  from,  obtained  by  lavage,  rhabditea 

found  in,  378 

—  larvae  of  Gastrophilus  inhabiting,  599 

—  trichomonads  in,   55 

—  Tristrongylus   instabilis   in,   435 

—  wall,  fibrous  thickenings  in,  produced  by 

species  of  Gnathostoma,  385 
Stomoxys,   characters,   609 

—  differentiation  of  Lyperosia  from,  610 

—  disease  carrier,  603 

—  species  of,   610 

—  calcitrans  (stinging  or  stable  fly),  609 
diseases  transmitted  by,  610 

ova,  larval  and  pupal  stages,  609 

transmission       of       epidemic       polio- 
myelitis by,  612 
Stools,  larvae  of  Blaps  mortisaga  in,  542 

—  method    of    discovering    head    of    tape- 

worms in,  674 
StreblidcB   (bat  parasites),  611 
Strepsiptera,  characters,  531 
Strong    and    Musgrave,    species    of    amoebae 

distinguished  by,  31 


INDEX 


891 


Strung ylidce,   375,  432 

—  free  life  of  young  stages,  20 
Strongyloides,    European,    free-living    gene- 
ration generally  absent  in,  383 

—  fulleborni,  384 

—  intestinalis ,     geogra2)hieal     distribution, 

384 
■ —  larvae  of,  cultivation,  474 
— -  life-history  of,  19 

—  morphology,   379 

—  stercoralis ,  free-living  form,  morphology 

of,    381 

generation,  female,  382  ■ 

habitat  in  body,   755 

—  —  heterogony   of,   381 

infection  by,  diagnosis,  675,  676 

diarrhoea  associated  with,   381 

expulsive  treatment,  675 

— pathological   significance,  674 

—  prophylaxis   against,   675 

symptoms,  674,  675 

larva  from  fresh  human  faeces,  382 

—  mature  filariform,  383 

mode  of  development,  373 

occurrence   in  man,   384 

—  —  parasitic  generation,  morphology,  381 
ova,  381,  382 

synonyms,   380 

and  Ancijlo stoma  duodenale,  larvae  of, 

diiit'erences  between,  451 

—  synonyms,   379 

—  toxic  action   of,  651 

—  vivipara,   384 
Strongylopkiimata,  208 

Stuelp,      amaurosis     following     male     fern 

poisoning,   671 
Stuertz,     chyluria    following    infection    by 

Eustrongylus   gigas,   682 
Stylorhynchus,  host  of,  135 

—  oblongatus,    gametes    of,    morphological 

differentiation,  133,  134 
Stypticin  in  bilharziasis,  643 
Sublimate,     corrosive,     saturated     aqueous, 

fixation  of  cestodes  by,  472 
solutions,  fixation  by,  748 

—  solution   in   crab   louse   infection,   712 
injection     in     expulsion     of     Guinea 

worm,   676 
into  cutaneous  and  muscular  cysti- 

cerci,  663 
Suckers  of  Cestoda,  289 
Sucking  worms,  see  Tr-etnatoda 
Suctoria,  29,   198 

—  characters  and  habitat,  29 

Sulphur,    flowers    of,    projjhylactic    against 
clothes  lice,  616 

—  preparations,  application  in  scabies,  706 
Sump    bunches,    skin    affection    set    up    by 

t  penetration    of    larvae    of    Ancylostoma 

duodenale,  455 
Surra,  animals  among  which  prevailing,  95 

—  causal   agent  of,   95 

—  geographical  distribution,  95 

—  transmission  by  Chrysops,  601 

by  Stomoxys,  96,  610 

by  Tabanus  sp.,  96 

56 


I  Surra,  trypanosomes  in  blood  of  horses 
with,  67 

Swallow  bug,  see  Cimcx  hirundinis 

Swammerdam,  discoveries  of  origin  of  para- 
sites, 10 

Swamps,  drainage  of,  in  prevention  of 
malaria,  636 

Sweden,  ox  Avarble  fly  (Hypodernia  bovis) 
attacking  man   in,  596 

Swellengrebel  and  Strickland,  on  Trypano- 
soma  lewisi,  92 

Symbiosis,   6 

Symmers,  bilharziasis  of  lung,  642,  643 

Symphoromyia,  characters,  603 

Syngamece,  characters,  459 

Syngamus,  459 

—  habitat  and  hosts  of  species,  459 

—  kingi,  habitat  and  host,  460 
morphology,   459,   460 

—  trachcalis,  bursa  of,  461 

Syphilis,  inoculation  with,  j^roducing  no 
immunity  to  yaws,   128 

—  non-immunity   to,    produced   by   inocula- 

tion with  yaws,  128 

—  parasite  of,   114,   124,   125,  632 

—  tertiary  eruptions  of,  Treponemata  diffi- 

cult to  find  in,  125 

—  treatment,   632 

Syrphidoz,    rat-tailed    larvae    of,    characters 

and  habitat,  583,  584 
^J^J^J  of   gregarines,    132 
Szerlicky,    case    of    intertrigo    set    up    by 

Oxyuris  vermicularis,  696 


Tabanid^  (gdd  flies),  characters,  600,  601 

—  diseases  transmitted  by,  601 

—  larvae,   600 

—  method  of  destruction,   601 

—  ova,   600 

—  pupae,   600 

Tabanus,  species  of,  transmitting  trypano- 
somes, 96,  601 

—  bovinus    (ox    gad    fly),    601 
Taenia,  331 

—  africana,  mature  segment  of,  342 
morphology,   342 

oncosphere   of,   299 

—  —  proglottis  and  head  of,  343 

—  bremneri,    morphology,    337 

—  capensis,  339 

—  coenurus,  nervous  system,  head  and  part 

of  neck  showing,  291 

—  confusa,    mature    and    gravid    segments, 

344 
morphology,  343 

—  crassicollis,     anatomy     of,     longitudinal 

section   showing,   290 

cysticercus  of,  338 

host   of,   6 

—  derivation  of  name   (footnote),  331 

—  echinococcus,  hooklets  of,  355,  359 
hosts  of,  345 

morphology,  344 

organs  of,  345 


892 


tHE   ANIMAL    PARASITES    OE   MAN 


Tcenia  echinococcus,  percentage  of  dogs 
infected  with,  in  various  cities  and 
countries   (footnote),  345 

—  —  rearing  of,  in  dog,  356 
synonyms,   344- 

see  also  Echinococcus 

—  expulsion  of,  resulting  in  cure  of  chorea, 

648 

—  extracts      of,      experimental      injection, 

effects,  648 

—  lata  (Dibothriocephalus  latus),  supposed 

origin  of,  11 

—  lophosorna,   339 

—  marginata,  337 

cysticercus  of,  338 

hooks  of,  338 

hosts  of,  338 

—  oncospheres      of      species      of,      animals 

selected  as  hosts  for  development,  299 

migration   from   intestine   through 

blood-vessels  to   liver,   302 

—  saginata,  cysticercus  of,  340 

expulsion  of,  best  method  for,  669,  67^ 

—  —  frequency  in  man,  341 

• genitalia,  proglottis  showing,  293 

geographical   distribution,  341 

habitat  in  man,  667 

host   of,   6 

malformations,   339 

morphology,  339 

parasitic  in  man  in  association  with 

other  tapeworms,  667 
proglottids    of,    feeding    experiments 

with,  340 

prophylaxis   against,   668 

race   incidence   of   infection,   340 

symptoms  produced  by   infection   by, 

667,  668 

synonyms,  338 

uterine  e^g,   298 

—  serrata,   cysticercus  of,  338 
hooks  of,  338 

host  of,  338 

—  solium,   339 
carriers  of,   335 

diagnosis  of  presence  in  body,  662 

Dipylidium    caninum    confused    with, 

680 

expulsion      from      body,      effect      on 

anaemia,  648 

measures  for,  must  be  thorough, 

336 

geographical    distribution,   334 

corresponds  with  that  of  domes- 
tic pig,  334 

■ habitat  in  body  of  man,  662 

head  of,  332 

host  of,  6 

in  man,  mode  of  infection,  335 

larval   infection,   662 

see  also  Cysticercus  celluloses 

malformations  of,  332 

modes  of  transmission,  336 

morphology,  331 

parasitic  association  with  Dibothrio- 
cephalus latus,  658 


Tcenia  solium,  proglottids,  332 

prophylaxis   against,   668 

symptoms   produced   by    infection    by, 

667,  668 
synonyms,   331 

—  species  of,  respective  times  required  for 

development  of  cysticercus  from  date  of 

infection,    304 
various,  respective  time  required  for 

growth,   306 
Taeniae,   development  of,   14 

—  infection  by,  treatment,  symptomatic,  669 

—  nervous  system  of,  290 

—  oncospheres  of,  14 

—  species  of,  in  relation  to  cystic  forms,  16 
Tceniidce,   331 

—  egg-shell  substance,  297 

—  eggs  of,  297 

—  morphology,  309 

—  oncospheres    of,    development    of    cysti- 

cerci   from,  303 

—  rostellum,   289 

of,   ring   encircling,   291,   292 

Taeniol,    administration   in    ancylostomiasis, 
686 

—  effects  of,  672 
Taeniorhynchus,  576,   577 

—  africana,  577 

—  annulipes,  577 

—  australiensis ,  577 

—  characters,  564 

—  major,  577 

—  ova   of,   557,   558,   577 

—  titillans,  577 

carrier  of  larvae  of  Filaria  bancrofti, 

577 

—  uniformis ,  bll 

carrier  of  larvae  of  Filaria  bancrofti, 

577 
Tallqvist,       experimental       bothriocephalus 

anaemia,  646 
Tamne  or  thimni  of  Kabyles,   598 
Taniguchi,  paragonimiasis  of  brain,  639 
Tapeworms,   adult,   length   of  life,   307 

—  biology,  307 

—  caudal  vesicle,  300 

—  cysticerci  experimentally  reared  from,  15 

—  development  of,  297 
embryonal,   298 

—  embryophore,  298 

—  experimental  rearing  of,   15 
from   cysticerci,   15 

—  expulsion  by  preliminary  aperients,  669 
by  vermifuges,  669 

—  found    in    association    with    other    intes- 

tinal parasites,  667 

—  individuality  of,  early  researches  as  to, 

283 

—  infection     by,     symptomatic     treatment, 

669 

—  injury  inflicted  by,  depends  on  number 

in   host,   9 

—  larvae    of,    sexual    maturity    must    take 
!      place   in   terminal   host,   304 

—  larval  stages,  development,  298-301 
modes  of,  300 


INDEX 


893 


Tapeworms,  metamorphosis  of,  15 
of  larva  into,  305 

—  method  of  discovering-  head  in  stools,  674 

—  oncospheres  (embryos)  of,  298,  299 
transformation    into    bladder    worms, 

303 

—  origin  of,  discovery,  11 

early    researches    as    to,    283 

—  ova  of,  297 

consistency,  297 

—  plerocercoid  of,  300 

—  scolex  of,  300,  303,  304 

Tar  and  sapo  viridis,   application  in  creep- 
ing disease,  732 
Tarsonemidce,  characters  of,  488 
Tarsonemus  intectus,  489 

—  uncinatiis ,  489 

Tartar  emetic  in  espundia,  629 

in   Indian   kala-azar,   626 

in   infantile   kala-azar,   627 

in  Oriental  sore,  628 

in  sleeping  sickness,  622 

Taschenberg,  Silvanus  surinamensis ,  542 
Taylor,  treatment  of  bronchial  spirochaeto- 

sis,  633 
Technique,       protozoological,       fixed       and 

stained  material,  747 

fresh   material,    745 

notes    on,    745-752 

Teeth,  carious,  spirochaete  associated  with, 

122 
Teichomyza  fusca,  larvae  of,  habitat,  584 
Teissier,  mercury  in  expulsion  of  Strongy- 

loides  stercoralis,  675 
Telosporidia,  28,   129 

—  characters,    28,    129 

Temnocephalid(B,  habitat  and  habits  of,  20 
Terebinthine   oil    in    chyluria   from   Filaria 

bancrofti  infection,  677 
Ternidens,    characters,   439 

—  deminutus,  439,  440 

habitat,  441 

Tersesthes,  581 

Testis,  enlarged,  in  filariasis,  401 

—  of  Ancylostoma  duodenale,  449 
Tetramitus,  57 

—  and  Chilomastix,  differential  characters, 

735,  736 

—  how  differing  from  Trichomonas,  57 

—  mesnili,  causal  agent  of  colitis,  57 
Fanapapea  intestinalis  identical  with, 

57 
habitat,  57 

—  —  synonyms,  57 

Tetranychidoe    (spinning    mites),    characters 

of,  488 
Tetranychus,   488 

—  molestissirnmus,    geographical     distribu- 

tion, 488 
itching  produced  by,  488 

—  telarius,  var.   russeolus,  effects  produced 

by,  488 
Tetratrichomonas,   53    (footnote),    734 
Texas  fever  in  cattle,  carriers  of,  177,  494 

causal  agent,  173,  177 

Theiler,  178,  180 


Theileria,  174,  178 

—  annulata,  180  , 

—  characters  of,  174 

—  mutans,  180 

—  parva,  178,   179 

agents  of  transmission,  179 

Koch's  blue  bodies  in,   179 

life-cycle  in  tick,  179 

morphology  of,  178 

pathogenic  agent  of  East  Coast  fever 

in  cattle,   174,   178 

—  stordii,  180 

Thelohan  on  Myxosporidia,  182,  183 

Thelohania  contejeani,  186 

Theobald,     F.      V.,     Arthropoda      (jointed 

limbed  animals),  483 
Theobaldia,  575 

—  annulata,  bite  of,  575 

characters,  575 

domestic  form,  575 

geographical  distribution,  575 

larvae  of,  habitat,  575 

—  characters,  564 

—  spathipalpis ,  bite  of,  575 
characters,  575 

geographical  distribution,  575 

Theobaldinella,  575 

Thiarsol  in  infantile  kala-azar,  627 

Thiopinol,    application    in    scabies,    706 

Thomas,  W.,  introduction  of  atoxyl  in  try- 
panosomiasis,  622 

Thomer,  treatment  of  crab  louse  infection, 
712 

Thomson,  D.,  sites  of  development  of 
crescents  of  tertian  malignant  parasite, 
169 

see  also  Ross,  Sir  R.,  and  Thomson, 

D. 

Thomson,  J.  D.,  researches  on  Trypanosoma 
lewisi,  89,  90,  92 

Thomson,  J.  G.,  and  Fantham,  cultivation 
of  Babesia  (Piroplasma)  canis  by  Bass's 
method,  172 

nuclear     phenomena     of     Babesia 

canis   in   cultures,   176 

see  also  Fantham  and  Thomson,  J.  G. 

Thomson,  J.  G.,  and  Sinton,  culture  of 
Trypanosoma  rhodesiense,  82,  83 

culture    of    trypanosome    forms    of 

T.  gambiense,  76 

medium  employed  by,   for  growth 

of  Trypanosoma  gambiense  and  T. 
rhodesiense,  745 

and  Thomson,  D.,  methods  of  cultiva- 
tion of  malarial  parasites,  171,  172 

number  of  merozoites  of  malig- 
nant tertian  parasite,  168 

spirochaetes    in    alimentary    tract, 

741 

Thornhill,  toxic  symptoms  following  thymol 
administration,  686 

"  Thymni  "   or  tamne  of  Kabyles,   598,  725 

Thymol,  administration  of,  in  expulsion  of 
ancylostomes,  685,  686 

mode  of,  685,  686 

of  ascarides,  694 


894 


THE   ANIMAL   PARASITES    OF   MAN 


Thymol,  administration  of,  in  expulsion  of 

Oxynridop,  697 

of  Strongyloides  siercoralis,  075 

in  flapfellate  dysentery,  624 

in  Trichiiris  trichiiira  infection,  679, 

680 
followed  by  benzene  enemata, 

680 
■ toxic  symptoms  following,  686 

—  enemata  in  arrest  of  trichinosis,  681 

in  expulsion  of  ascarides,  694 

Thymoluria,  686 

Thymotol,      administration      in      ancylosto- 

miasis,  686 
Thyroiditis,   parasitic,   87 

—  see  also  Trypanosomiasis ,  Brazilian 
Thysanoptera,  characters,  531 

Tick,  stag-es  of  life-cycle  of  Babesia  canis 
and  B.  hovis  in,  176,  177 

—  bites,  paralysis  due  to,  613 

—  or   relapsing-   fever,    116,    630 

African,   carrier  of,    116,  496,  630 

• importation    into    Persia,    613 

pathogenic  agent,  116 

—  paralysis,  cause  of,  504 

Ticks,  transmission  of  piroplasmosis  by,  from 
recovered  to  uninfected  animals,  178 

Tiger,  host  of  Paragonimus  westermannii; 
250 

Tinea  rotunda,  see  Ascaris  lumhricoides 

Toad,  rectum  and  urinary  bladder  of, 
Opalina  parasitic  in,  207 

Todd,  on  leucocytogregarines  in  birds,  154 

—  tick  paralysis,  613 

—  see  also  Button  and  Todd 
Tommasi-Crudeli,       early       researches       on 

malaria,  156 

Tomsk,  Opisthorchis  felineus,  human  para- 
site most  frequently  found  at  autopsies 
at,  253 

Tongue,  cysticercus  of,  663 

Townsend,  Simulium  occidentaJis ,  579 

Toxascaris,   characters,   465 

—  limhata,   morphology,   466 
ovum  of,  466 

synonyms,   466 

Toxoplasma,   112 

—  hosts  of,  113 

—  pyrogenes.     association     with     splenome- 

galy, 113 
Toxorhynchites,   characters,   563,   570 
Trachea,  ascarides  invading,  691 
Trachoma  bodies  in  infected  epithelial  cells 
of  conjunctiva.  209   (fig.   119) 

so-called,   cultivation,   210 

Trematoda.  endoparasitic  life  spent  in  in- 
termediate and  final  host,  18 

—  relation  to   Turhellaria,   19 
Trematodes   (sucking  worms  or  flukes),  212 

—  ao-e   attained   by,   230 

—  alimentary  canal,  217 

—  asexual    generations,   224,   225 

—  cercariae   (larval   stages),   225,  227,  228 

—  cirrus  sac,  221 

—  copulation    iii,   222 
-.  —  cross,   22? 


Trematodes,  development,  12,  222 

embryonic   and   post-embryonic,   224 

final,  conditions  necessary  for,  225 

—  developmental   cycle,  229 

—  digenetic,     adult,     animals    harbouring, 

how   infected,   226 

development,  224,  226 

miracidia  of,  226 

—  endoparasitic,  biology  of,   229 
hosts  and  habitat  of,  229 

—  excretory  bladder,  219 
system,   219 

terminal  flame  cell,  219 

—  food  of,  218 

—  found  in  man,  classification,  230 

—  genital  pore,  222 

—  intestine  of,  variation   in,  217 

—  investing  layer  of,  213 

—  Laurer's  canal  of,  221,  222 

—  metraterm  of,   221,  222 

—  miracidia  of,  223,  224 

—  morphology  of,  212 

-  movements  of,  216 

-  muscular  system   of,   214 

-  nervous  system  of,  216 

—  organs  of  sense,  216 

—  origin  of,  12 

of  parasitism  in,  20 

—  ova  of,  deposition,  223 
formation,  223 

—  parenchyma  of,  213 
muscles   of,   214 

—  redise  of,  225,  226,  227,  228 

—  salivary  glands,  217 

—  sexual   organs,  220 

deviation     from     typical     position 

(footnote),  222 

female,  220,  221 

male,   220 

—  shell  gland  secretion  in,  223 

—  sporocyst  of,   225,   227 

—  suckers  of,  213,  214 

—  and  turbellaria,  genetic  relationship  be- 

tween,  20 
Treponema,    114,    115,    123 

—  calliqyrum,   126 

—  cultivation      of      species      from      human 

mouth,   128,   741 

—  macro  d  entium ,   128 

—  microdentium ,  128 

—  morphology,  124 

—  mucosum,   128 

—  pallidum,  114 

causal  agent  of  syphilis,  124 

^  — -  cultivation  of,  method,  125 

difficult  to  find  in  tertiary  eruptions 

of  syphilis.   125 

granule  fcrmation,   124,  125,  127 

morphological    and    pathogenic   varia- 
tions, 126 

morphology,   124,   125 

Svirochoeta        refringens        associated 

with,   122 
- — •  —  synonyms,    124 

—  pertenue,   ciiltivation,   128 
granule   formation,   127 


INDEX 


895 


Treponema  pertenue,  mode  of  infection,  128 

morphology,    127 

pathogenic  agent  of  yaws,  114,  127 

reasons  for  considering  specific  cause 

of  yaws,  128 

—  species    of,    association    with    pyorrhoea 

alveolaris,  128 
Treutler,    filaria    associated    with    phthisis, 
408 

—  parasite,    probably    liver-fluke,    in    vein, 

243 
Triaenophorus,      excretory      vessels,      island 
formation,   292 

—  plerocercoid  of,  300 

Triatoma  megista,  discovery  of  Trypano- 
soma cruzi  in,  83,  84 

jihases  of  development  of  Trypano- 
soma cruzi  in,  87 

preventive  measures   against,   623 

Triboulet,  Ascaris  infection  in  relation  to 
appendicitis,  653 

Trichina  spiralis,  423 

Trichinella,  421 

—  development   in    definite   host,    18 

—  spiralis,  421 

development   of,   373 

history  of,  423,  424 

geographical      distribution      not      in 

correspondence  with  occurrence  of 
trichinosis  in  man,  427,  428 

hosts  of,  6 

in      man,      percentage      of      invasion 

according  to  nationalities  determined 
by  post-mortem  examination,  428 

infection   by,   680 

distribution  in  body  after,  424 

— see  also  Trichinosis 

invasion   and  encystment  in  muscles, 

424,   425 

mammals  in  which  developed  experi- 
mentally, 421 

infected  by,  in  order  of  frequency, 

421 

inhabited  by,  421 

morphology,  421 

normal  hosts  of,  427 

—  —  symptoms  produced  by,  in  periods  of 

invasion,  dissemination  and  encystment, 
424,  425 

viviparoiis  nematode,  371 

Trichinellse,  development  in  encysted  con- 
dition, 427 

—  encysted,    in    man    and    other    mammals, 

early  observations  of,  423 

—  fatal  case  of  infection  by,  423 
- —  feeding  experiments  with,  423 
Trichinellida?,  419 

—  characters,  375 
Trichinellince,  421 

Trichinosis,  amount  of  prevalence  in  North 
America,  428 

—  diagnosis,  681 

-by  blood  examination,  681 

—  epidemics  of,  423 
in   Germany,  423,   429 

r —  geographical  distribution,  428 


Trichinosis  in  man,  geographical  distribu- 
tion of  Trichinella  spiralis  not  in  cor- 
respondence with  occurrence  of,  427,  428 

—  prophylaxis  against,  429,  431 

—  symptoms   of,   424,   425,   680 

—  treatment,  before  and  after  development, 

681 
Trichocephali  in  appendix,  655 
Trichocephalus  anaemia,  651 

—  infection   by,  effects  of,  651 
in   relation   to   appendicitis,   653 

—  lacks  intermediate  host,  21 
Trichomonads,  habitat  in  body,  55,  735 

—  question    of   cysts   of,    56 
Trichomonas,  52 

—  characters  of,  52 

—  diarrhoea  due  to,   57,  624,   734 

—  from  gut  and  caecum  of  rat,  735 

—  hominis  same  as  T.  intestinalis,  54 

—  intestinalis,   45,    54 

—  — •  axostyle   of,   55 

characters  of,  55 

flagella   of,   55 

relation  to  T.  vaginalis,  54 

spherical    contracted    forms    in    mice, 

56 
transmission,  modes  of,  56 

—  points  of  difference  of  Tetramitus  from,  57 

—  regions  of  body  other  than  intestine  in 

which  found,  55,  56 

—  vaginalis,  52,  760 

characters  of,  52,  53 

flagella  of,  53 

nucleus  of,   53 

—  —  presence  in  urethra  of  male,  53 
Trichomoniasis,    human,    recent    researches 

in,   734 

—  oral,  treatment,  625 

—  vaginal,  treatment,  625 
Trichopalpus,   603 

—  larvae,  characters  and  habitat,  603 

—  ohscurus,   603 
Trichoptera,  characters,  531 
Trichosoma  crassicaudum,  female  parasitic,  4 

habitat  of,  4 

Trichostrongylince ,   characters   and  habitat, 

433 
Trichostrongylus,  morphology,  434 

—  instabilis,   habitat,  435 
hosts  of,  435 

^  in   man,    cases   recorded,    435 

morphology,  434 

—  probolurus ,  habitat,  435 
hosts  of,  435 

morphology,  435 

—  vitrinus,  hosts  of,  436 

morphology,  435,  436 

Trichotrachelida-,  oesophagus  of,  363 

—  unicellular   cutaneous   glands   of,   361 
Trichurince,  419 

Trichuris,   morphology,    419 

—  alcocki,  421 

—  cameli,  421 

—  campanula,  421 

—  crenata,   421 

infection  with,  420 


896 


THE   ANIMAL   PARASITES    OF   MAN 


Trichuris  depressiuscula,  420,  421 
infection   with,  420 

—  discolor,  421 

—  giraffcB,   421 

—  globidosa,  421 

—  nodosus,  421 

—  avis,  421 

infection  with,  420 

—  trichiura,  habitat  in  man,  420 

infection  with,  sources,  679 

symptoms,   679 

treatment,   679 

mammals  inhabited  by,  421 

mode  of  attachment  to  wall  of  in- 
testine, 679 

morphology,  419 

ova,   development   of,   420 

embryo-containing,  420 

infection  by,  420 

parasitic  in  large  intestine,  678 

percentage  found  at  autopsies,  420 

synonyms,  419 

—  unquiculata,  421 

Trinidad,  mosquito  worm  in,  598 
Triodontophorus,      bursal      formula      (foot- 
note),  439 
Troglotremidcs,  249 

—  morphology,  232 
Trombidiidas,  characters,  485 
Trombidium,  485 

—  fuliginosum,  486 

—  gymnopterosum,  486 

—  serraticeps,   486 

—  tlalsahuate,  skin  affections  set  up  by,  486 
Trophozoites  of  Coccidia,  140,  143 

—  of  Entamoeha  tetragena,  39,  40 

—  of  gregarines,    132 

—  of  malarial  parasites,   159 

—  of  Microsporidia,  185 

—  of  Myxosporidia,   182 

Tropical  Diseases  Bureau  Bulletin,  founda- 
tion   of,   69 

Tropical  sore,  see  Oriental  sore 

Trouessart,  Histioqaster  (entomophagus  ?) 
spermaticus ,   515 

Trypan-blue  treatment  of  piroplasmosis,  178 

dosage  for  dogs,  horses  and  cattle,  178 

Trypanophis,   63 

Trvpanoplasma,   characters  of,  63 

—  hosts  of,  63 

T:  ypanoplasms  in  iish,  68 
Trypanosoma,  67 

—  americanum,    69 

—  hoylei,   99 

experimental  infection  with,  99 

host   of,   99 

—  hriicei.  93,  94 

and  T .  rhodesiense,  question  of  dis- 
til-ction    or    identity,   80,   83,    94 

blepharoplastless  strains,  101,  737 

cause  of  nagana  ("tsetse-fly  disease),  93 

development  in  Glossina  morsitans,  94 

drug  resistance  of,  101 

innocuous  to  big  ffame,  70 

morphology  and  life-history  in  verte- 
brate host,  94 


Trypanosoma  hrucei,  nucleus,  blepharoplast 
and  flagellum  of,  70 

posterior  nuclei  in,  83 

strain    from    Uganda,    95 

from    Zululand,    94,    95 

—  caprce,  monomorphic,  100 

—  cazalboui,  causal  agent  of  "  souma,"  100 
monomorphic,  100 

—  characters,  67 

—  congolense,  agents  of  transmission,  100 

cause  of  Gambia  horse  sickness,   100 

geographical   distribution,    100 

monomorphic,  100 

probable  synonyms,  100 

—  cruzi,  83 

crithidial   forms,   86 

culture,   87 

geographical  distribution,  83,  84 

hosts  of,  85,  86,  87 

in   foetus,  88 

invertebrate  host  of,  83,  84,  537 

life-history  in  invertebrate  host,  80 

in  vertebrate  host,  84 

modes       of       multiplication 

("  sexual "  and  asexual),  85,  86 

microgametes    and    macrogametes,    85 

morphology,  84 

possible  reservoir  of,  87 

schizogony  of,  84,  85,  86 

—  dimorphon,    100 

—  equi,   83,    98 

—  equinum,  cause  of  "  mal  de  caderas,"  96 
morphology,   96 

transmission  of,  97 

—  equiperdum,  97 

cause      of      "  dourine "      or     stallion 

disease,  97 

endotoxins  in,  98 

morphology,  98 

posterior  nuclei  in,  83 

progress    of    disease,    97 

—  evansi,  blepharoplastless  strains,  737 

causal  agent  of  surra,  95 

morphology,  95,  96 

possible  case  in  man,  96 

synonyms,  95 

transmission  of,  95 

experimental,  67 

variety  causing  "  mbori  "  in  drome- 
daries,  96 

—  fringillarum,  737 

—  gambiense,   68,   72 

cause  of  sleeping  sickness,  68,  605 

cultivation  of,  medium  used  for,  745 

cultures  of  trypan osome  forms  of,  76 

development  in   Glossina  palpalis,  74, 

75 

effect  of  serum  reactions  on,  80 

immunization  against,  does  not  pro- 
tect against  infection  by  T.  rhodesiense, 
80 

in    antelope,    76 

innocuous  to  big  game,  70 

invasion   of   salivary   glands   of    Glos- 
sina palpalis,  75 
latent  forms  of,  77 


INDEX 


897 


Trypanosoma   gambiense,  morphology,   72 

in  circulating  blood,  73 

serum  from  animals  infected  Avith,  no 

effect  on  T.  rhodesiense,  80 

trypanolytic   for,   80 

synonyms,  72 

- —  hippicum,  agents  transmitting,  99 

■ cause   of   "  murrina "   in   mules,   98 

morphology,   98 

—  lewisi,  crithidial  forms,  91 
development   in    rectum    of   rat 

flea,  91,  93 

inoculation  experiments,  90 

life-cycle  in  invertebrate  host,  90,  91 

in  vertebrate  host,  88,  89 

morphology,  88 

multiplication  rosettes,  71 

potential  pathogenicity,  737 

—  —  rosette   forms,   89,    90 

strain  of,  losing  resistance  to  arseno- 

phenyl-glycin,   how  effected,  93 

—  —  transference    from    blood    of    rat    to 

blood  of  snake,  102 
transmission  of,  88 

—  nanum,   100 

—  nigeriense,  76 

—  noctiicB,    69,    737 

—  pecaudi,  95 

causal   agent  of   baleri   in   sheep   and 

equines,  95 
posterior  nuclei  in,  83 

—  pecorum,   100 

—  rhodesiense,   69,   76 

and  T .  brucei,  question  of  distinction 

or  identity,  80,  83,  94 

animal  reactions,  78 

cause  of  Rhodesiau  sleeping  sickness, 

69,  76,  605 

cultivation  of,  83 

medium  used  for,  745 

developmental  cycle  in  Glossina  mor- 

sitans,  81 

effect  of  serum  reactions  on,  80 

Glossina  morsitans   transmitting,   608 

immunization    against    T.    gambiense 

does  prevent  infection  by,  experiments 

proving,  80 

latent  or  resting  forms  of,  77,  78 

morphology,  76,  77 

non-pathogenic  to  antelopes,  70 

partial  immunity  against,  81 

pathogenic    to    man    and    laboratory 

animals,  70 

posterior  nuclei  in,  83 

reservoir  of,'  81 

resistant  to  atoxyl,  78 

—  —  serum  from  animals  infected  with  T. 

gambiense  has  no  effect  on,  80 

transmission  of,  69,  81 

climatic    factors    affecting,    81 

virulence   of,   compared  with  that  of 

T.  gambiense,  78 

—  simioB,  virulent  to  monkeys  and  pigs,  100 

—  theileri,    98,    611 

—  —  geographical  distribution,  98 
morphology,  98 


Trypanosoma  ugandce,  95 

—  uniforme,  hosts  of,  101 
monomorphic,    101 

—  vivax,   fatal  to  cattle,  99 

—  —  monomorphic,  99 

transmission   of,    100 

Trypanosome,    animal,    infection    of    human 

being  with,  96 

—  diseases  spread  by  Glossina,   603 

—  human,   68,   69 

■  artificial      infection      of      species      of 

Glossina  with,  605 

—  infections,  Liverpool  School  of  Tropical 

Medicine    Expedition    sent    to    investi- 
gate,  68 
Trypanosomes,  adaptation  of,  101 

—  artificial   cultivation,   69 

—  blepharoplastless,   101,   737 

—  classification,  71,  72 

—  deleterious  or  fatal  to  domestic  animals, 

69 

—  general    note    on    development    in    Glos- 

sina, 101 

—  hosts  of,   67,  68,  69 

—  immunity  to,  in  antelope,  69 

—  in  blood,  cultures  aid  in  detection  of,  69 
~   —  cyclical   variation,   78 

—  —  daily  number  from  case  of  Rhodesian 

sleeping  sickness,  79 

method  of  determining  number,  748 

multiplication,  71 

periodicity,   69 

seasonal  variation,  69 

—  in     cerebrospinal     fluid     from     cases     of 

sleeping  sickness,  68 

—  latent    forms,    non-flagellate,    from    in- 

ternal organs  of  vertebrates,  73,  74,  77 

—  monomorphic,  99 

—  morphology  of,  70 

—  nuclei  of,  70 

—  pathogenic  to  man  and  domestic  animals, 

70 

—  percentage  of  fleas  fed  on  infected  rat 

becoming  infected   with,   93 

—  polymorphism,  72 

—  posterior  nuclei  in,  83 

—  resting   stages,   72 

—  transmission  from  one  vertebrate  host  to 

another,  72 

—  transmissive  stage  in  vertebrates,  737 

—  transmitted  experimentally  by  Stomoxys, 

610 

—  undulating  membrane,  71 
Trypanosomiasis,      African,      see     Sleeping 

sickness 

—  Brazilian,   acute,  87 

chronic,   varieties   of,   87,   88 

clinical  features,  87 

hereditary  transmission,  88 

histopathology,  88 

suggested   treatment,   623 

synonyms,  87 

—  cryptic,  69 
TrypanosomidcB,  61 

—  characters,  66 

—  genera  of,  67 


898 


THE    ANIMAL    PARASITES    OF    IMAN 


Tryposafrol,      producing      blepharoplastless 

trypanosomes,    737 
Tsetse-fly,  see  Glossina  morsitans 

—  disease,  see  Nagana 
Tuberculosis,  see  Cestode  tuberculosis 
"Tuft-like"    or    "phagocytic"    organs    of 

nematodes,  362 

Tumours,  subcutaneous,  associated  with  in- 
vasion by  Onchocerca  volvulus,  418 

Turbellaria,  parasitic,  2 

—  relation  of  Trematoda  and  Cestoda  to,  19 

—  and  trematodes,  genetic  relationship  be- 

tween, 20 
Turkeys,  blackhead  in,  145 
Turpentine  in  flagellate  diarrhoea,  624 

—  in  nasal  myiasis,  719 

—  oil   of,   in   bilharziasis,   643 
Tydens  molestus,  habitat,  491 

host-tormenting,  491 

Tylenchus  putrefaciens ,  379 

Typhlitis,    association   of    Oxyuris   vermicu- 

laris   with,  467 
Typhlocoelum  flavum,  progeny  of,  discovery, 

12 
Typhoid   fever,   helminthes   as   predisposing 

factor  of,  657 

peculiar   fever   resembling,   613 

spread   by   house-fly,   586 

symptoms  of,  in  lumbricosis,  650 

—  vaccine  in  bilharziasis,  644 

Typhus,    possibly    due    to    a    chlamydozoon, 

207 
Tyroglyphi,    differentiation    of    Glyciphagi 

from,  513 
TyroglyphidcB,  characters,  511 

—  habitat  and  food  of,  511 
Tyroglyphus  longior,  512 

characters  of,   512 

habitat,  512 

—  minor,    var.    castellani,    cause    of    copra 

itch,  513 

—  siro,  512 

—  —  characters  of,  511 


U. 


Uganda,     strain     of     Trypanosoma     brucei 
from,  95 

—  syphilis   in,   treatment,    632 
TJhlenhuth  (and  others),  endotoxins  in  Try- 

panosoma  equiperdum,  98 
Ulcers  arising  from  clothes  louse  infection, 
711 

—  and  boils  due  to  invasion  by  Cordylobia 

anthropophaga,   592 

—  examination  for  protozoa,  746 

—  production  by  species  of  Enyaliopsis ,  542 
Ulcus  tropicum,  agent  of,   122 
Umbilicus,  ascarides  escaping  from,  656 
Unger,  treatment  of  oxyuriases,  697 
Uranotsenia,  characters,  565 

Urethra,    fistulae    of,    arising    from    bilhar- 
ziasis, 642 
treatment,  644 

—  larvae  of  Homalomyia  canicularis  found 

in,  585 


Urethra,  maggots  passed  from,   728 

—  male,    presence    of    Trichomonas    vagina- 

lis  in,   53 
Urinary  apparatus,  symptoms  of  bilharzia- 
sis mainly  centred  in,  641 

—  passages,  invasion  by  ascarides,  692 
Urine,  amoebae  found  in,  45,  46 

—  human,  aphides  said  to  have  been  passed 

in    (footnote),   532 

—  occurrence  of  Anguillula  aceti  in,  379 

—  presence    of    Nephrophages    sanguinarius 

in,   490 

—  preservation  of  ova  of  flukes  in,  472 
Urosporidium,   194 

—  fuliginosum,   195 
Urotropine  in  bilharziasis,  643 
Urticaria,   echinococcus   cysts   causing,   651, 

652 

—  set  up  by  Leptus  autumnalis,  702 
Uterus,    cervix,    polypoid   tumour    of,    with 

Schistosoma   infection,   643 
Uzara   in   flasrellate   diarrhoea,   625 


Vaccine    and    emetine    treatment    combined 

in   pyorrhoea  alveolaris,   620 
Vaccinia,  cell  inclusions  in,  207,  208 
Vagina  atrophied  in  Acole'inoB,  297 

—  presence  of  Bhabditis  pellio  in,  377 
Vaginitis,  acute,  due  to  Schistosoma  infec- 
tion, 643 

Vanilli.smus,  so-called,  cause  of,  512 
Varicose  glands  in  filariasis,  402 
Variola,   cell   inclusions   in,   207,   208 
Vegetable  food,  raw,   avoidance  of,  in   pro- 
phylaxis against  Oxyuriasis,  697 

—  matter,    decomjoosing,    Tyroglyphida;    in, 

511 
larvae     of     Homalomyia     canicularis 

found  in,  585 
Veins,  liver-flukes  found  in,  243 
Vena  cava  and  portal  vein,  communication 

between,   how  formed,   272 
Verallina,  characters,  565 
Vermifuges,  669-675 
Vertebrates,   entamoebse   of,   34 

—  experimental      introduction      of      insect 

flagellates  into,   104,   112,  737,  738 

—  internal  organs  of,  latent  forms  of  try- 

panosomes   from,   73,   74 

—  multiplication  of  trypanosomes  in  blood 

of,  71 

—  spirochaetes  in,  116,  122 

Vesicles,  formation  of,  in  creeping  disease, 
730 

Vesico-prostatic  plexus.  Schistosoma  haema- 
tobium in,  273,   274 

Vianna,  histopathology  of  Brazilian  try- 
panosomiasis,   88 

—  treatment  of  espundia,  629 

Viereck,   discovery  of  Entamceba  tetragena 

by,  38 
Vignolo-Lutari,  case  of  intertrigo  set  up  by 

Oxyuris   vermicularis,  696 


INDEX 


899 


Villot,  larvae  of  Goirliidoe,  479 

Vineg^ar,    Anguilliila   aceti   found   in,   379 

—  see   also  Sabadill  vinegar 

Virchow,    R.,    development    of    Trichinella 

spiralis,  423 
doubtful    case    of    human    coccidiosis, 

149 

Echinococcus  multilocularis ,  356 

Vital,  liver-fluke  in  vein,  243 

Vleming-kz's       mixture,        application        in 

scabies,   706 
Vog-t,   C,   on   the   Helminthes,   3 
Vomited  matter,  spirochaetes  in,  122 
Vorticella  in  faeces,  206 


W. 


Wagener,  von,  lesions  produced  by  Oxyuris 
vermicularis ,   695  ' 

—  life-history  of  Oxyuris  vermicularis ,  467 
Waldenburg-,    experimental    infection    vs^ith 

Coccidia,   136 
Walker     and     Sellards,     experiments     with 

dysenteric  amoebae,  618 
Walker,  E.  L.,  balantidiasis,  203 

on  Entamoeba  histolytica,  40  | 

prevention  and  treatment  of  balanti- 

dian    dysentery,    637  j 

Walker,  Norman,  treatment  of  scabies,  707    ' 
Walrus,  host  of  Dibothriocephalus  cordatus,    \ 

315  .  ' 

Walsh  and  Riley,  Rasahus   biguttatus,  540 

Beduvius  personatus ,  540 

Warble   uieo   {uesiriace),   liosts  of,  594  j 

Warburg,  extract  of  male  fern  in  expulsion    I 

of  ancylostomes,  687  i 

Wasielewski,     Hoemoproteus     (Halteridium)    \ 

danilewskyi,  var,  falconis,  152  [ 

Water,   eggs  of  mosquitoes  float  on,  559        ! 

—  filtered     and     boiled,     as     prophylactic 

ag-ainst  bilharziasis,   644 

—  infected,    avoidance    of,    in    prophylaxis 

against  Guinea  worm  infection,  676 

—  larvae  of  Stegomyia  fasciata  occur  in  all 

collections  of,  574 

—  mature  larvae  of  Ancylostoma  duodenale 

capable  of  living-  in,  454  j 

—  receptacles,    screening    against    mosqui-    i 

toes,  636  j 

—  stagnant,  mosquitoes  depositing  ova  in,    j 

553,  557 

—  transmission     of    trichomonad    infection    j 

by,  56,  624  j 

—  weeds    harbouring   mosquito   larvae,    des-    : 

truction  of,  636  I 

Watercress,   passage  of  larvae  of  Syrphidce 

into  human  beings  through  eating,  584 
Watsonius,  234 

—  watsoni,  234,  235 

—  —  diarrhoea  in  host  associated  with,  235 

female  organs,   235 

habitat,  235 

male  organs,  234 

morphology,  234 

ova,  235 

57 


Watsonius  watsoni,  synonyms,  234 
Weichselbaum,   intestinal   myiasis,   726,   727 
Weidman,   see  Smith  and    Weidman 
Welland,   Ascaris  sp.,  465 
Wellmann,  the  ochindundu,  541 
Wendelstadt     and     Fellmer,     trypanosomes, 

mutation   experiments   with,   102 
Wenyou,    C.    M.,    connection    of    Cimex    sp. 

with  Oriental  sore,  536 
possible   host   of   Leishmania   tropica, 

108 

on  Entamoeba  histolytica,  40,  41 

on  genus  Cercomonas,  736 

—  spherical    contracted    forms    of    Tricho- 

monas  intestinalis ,   56 

—  supposed    intermediate    host    of    parasite 

of  Bagdad  sore,  575 

—  Tetramitus   mesnili,   57 

—  transmission  experiments  with  Trypano- 

soma lewisi,  92,  93 
Werbitzki,    blepharoplastless   trypanosomes, 

101 
Wheler,  Dermacentor  reticulatus,  502 

—  length  of  life  of  Ixodes  plumbeus    (dog 

tick)   apart  from   host,  495 

—  life-history  of  Ixodes  reduvius,  494 
Whip  worm,  see  Trichuris  trichiura 
White    mice,     experimental    production     of 

disease  like  leishmaniasis  in,   103 

infection    with    Herpetomonas    cteno- 

cephali  and  H.  pattoni,  103 

—  scour  in  fowls,  causal  agent,  145 
Whitfield,   A.,    and    Hobday,    F.,   transmis- 
sion of  dog  mange  to  man,  523 

Whittles,  nematode  larvae  in  periosteum  of 
upper  jaw  in  case  of  gingivitis,  378 

Wiggins,   locust  injurious  to  man,  542 

Wijnhoff,  cases  of  amoebae  in  urine,  46 

Wild  game,  Trypanosoma  rhodesiense  pre- 
sent in,  81 

Wilkinson's  ointment,  application  in 
scabies,   706 

Williams,  Anna  W.,  culture  media  for 
amoebae,  743 

on  cultural  amoebae,  42 

Williams,  H.  XJ.,  invasion  of  human  beings 
by  Trichinella  according  to  nationali- 
ties,  428 

Wilms,  myiasis  oestrosa  dermatosa,   725 

Winogradoff,  post-mortem  discoveries  of 
Opisthorchis  felineus,  252,  253 

Wirsing,  mode  of  infection  of  intestinal 
myiasis,  727 

Wohlfahrt,  myiasis  cutanea  from  Sarco- 
phaga,  722 

Wolff,  treatment  of  cutaneous  and  muscu- 
lar cysticerci,  663 
Woodcock,    transmissive   phase   of   trypano- 
somes,  737 
Wood  tick,  see  Dermacentor  occidentalis 
Worm  abscesses,  formation  of,  9 

—  electuary      (Stork's)      in      expulsion      of 

ascarides,  692 

—  seed  oil  in  expulsion  of  ascarides,  694 

"  Wormlet "  burrowing  into  human  epider- 
mis, 599 


900 


THE   ANIMAL    PARASITES   OF   MAN 


Worms,   intestinal,   hereditary   transmission 

of,  former  belief  in,  11 
of     lower     animals     represent     young 

stages,   21 

spontaneous  generation,   belief  in,    12 

transmission  by  ova,  discovery  of,  11 

Wounds,   larvae   in,   movements   of,   723 
Wright,    Bhinosporidium    kinealyi,    197 
Wurtz  and  Cleri,  invasion  by  Loa  loa,  678 
Wyeomyia,  characters,  565 


Xenopsylla,   distinctive  characters,  545 

—  host     of     cysticercoids     of     Hymenolepis 

murina  and  H.  nana,  328 

—  brasiliensis,   547 

—  cheopis,  546 

carrier  of  plague  bacillus,  543,  547 

host  of  Trypanosoma  lewisi,  92 

Xeroform,   application  in   Demodex  follicu- 

lorum  canis  infection,  709 
Xyphorhyncus  firmus,  131 


Y. 


Yaws,  climatic  distribution,   632 

—  inoculation   with,   experimental,   127 
producing   no    immunity   to    syphilis, 

128 

—  non-immunity   to,    produced    by    inocula- 

tion with  syphilis,   128 

—  pathogenic  agent  of,   114,   127,   128,  632 

—  prophylaxis,  632 

—  species  of  Sarcophaga  concerned   in   dis- 

semination of,  590 

—  stages  of,  632 

—  treatment,  632 


Yellow  fever,  mosquito  carrier  of,  574 

Paraplasma    fiavigenum    said     to     be 

associated    with,    180 
transmission    by   Stegomyia,   555 

—  pigment    in    kidney    and    liver    cells    in 

ancylostomiasis,   647 
Yorke   and  Blacklock,   classification   of  try- 
jjanosomes,  72 

—  see  also  Blacklock  and  Yorke 

—  see   also  Stannus  and   Yorke 


Z. 


Zarniko,  case  of  Oxyuridoe  in  nose,  696 
Zeder,  special  class  of  cysticerci  established 

by,   282 
Zeller,   Echinococcus   multilocularis ,   356 
Zenker,  development  of  Trichinella  spiralis, 

423 

—  fatal    case    of   infection   by    Trichinellae, 

423 

—  Linguatula  serrata,  527 
Zenker's  solution,  749 

Ziemann,  infection  by  Loa  loa,  678 

—  varieties  or  sub-species  of  malignant  ter- 

tian  parasite,    167 
Zinn,  blood-stained  diarrhoea  from  Strongy- 
loides  stercoralis  infection,  674 

—  extract    of    male    fern    in    expulsion    of 

ancylostomes,   687 
Zooparasites,   1 
Zschokke,    experimental    infection    of    man 

with  Dibothriocephalus   latus,  312 

—  Rhinosporidium  in  horses,  197 
Zuelzer,  on  spirochsetes,  114,  741 
Zululand,     strain     of     Trypanosoma    brucei 

from,  94 
Ziirn,   case  of  transmission   of  infection    by 

Demodex  folliculorum  canis  to  man,  709 
Zygotes  of  Coccidia,  141,  144 

—  of   gregarines,    132,    133 


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LIBRARY,  UNIVERSITY  OF  CALIFORNIA,  DAVIS 

Book  Slip-Series  458 


U 71201 

Fantham,  H.  B. 

Animal  parasites 
of  man. 


QL757 

F3 


LIBRARY 

UNIVERSITY  OF   CALIFORNIA 

DAVIS 


